anatomy

Gheerbrant, E., 2009. Paleocene emergence of elephant relatives and the rapid radiation of African ungulates
43. Proc. Natl. Acad. Sci. U. S. A 106, 10717-10721.
Abstract: Elephants are the only living representatives of the Proboscidea, a formerly diverse mammalian order whose history began with the 55-million years (mys) old Phosphatherium. Reported here is the discovery from the early late Paleocene of Morocco, ca. 60 mys, of the oldest and most primitive elephant relative, Eritherium azzouzorum n.g., n.sp., which is one of the earliest known representatives of modern placental orders. This well supported stem proboscidean is extraordinarily primitive and condylarth-like. It provides the first dental evidence of a resemblance between the proboscideans and African ungulates (paenungulates) on the one hand and the louisinines and early macroscelideans on the other. Eritherium illustrates the origin of the elephant order at a previously unknown primitive stage among paenungulates and "ungulates." The primitive morphology of Eritherium suggests a recent and rapid paenungulate radiation after the Cretaceous-Tertiary boundary, probably favoured by early endemic African paleoecosystems. At a broader scale, Eritherium provides a new old calibration point of the placental tree and supports an explosive placental radiation. The Ouled Abdoun basin, which yields the oldest known African placentals, is a key locality for elucidating phylogeny and early evolution of paenungulates and other related endemic African lineages

Hakeem, A.Y., Sherwood, C.C., Bonar, C.J., Butti, C., Hof, P.R., Allman, J.M., 2009. Von Economo neurons in the elephant brain. Anat. Rec. (Hoboken. ) 292, 242-248.
Abstract: Von Economo neurons (VENs), previously found in humans, all of the great ape species, and four cetacean species, are also present in African and Indian elephants. The VENs in the elephant are primarily found in similar locations to those in the other species. They are most abundant in the frontoinsular cortex (area FI) and are also present at lower density in the anterior cingulate cortex. Additionally, they are found in a dorsolateral prefrontal area and less abundantly in the region of the frontal pole. The VEN morphology appears to have arisen independently in hominids, cetaceans, and elephants, and may reflect a specialization for the rapid transmission of crucial social information in very large brains

Jantou, V., Turmaine, M., West, G.D., Horton, M.A., McComb, D.W., 2009. Focused ion beam milling and ultramicrotomy of mineralised ivory dentine for analytical transmission electron microscopy
114. Micron. 40, 495-501.
Abstract: The use of focused ion beam (FIB) milling for preparation of sections of mineralised ivory dentine for transmission electron microscopy (TEM) is investigated. Ivory dentine is essentially composed of fibrillar type-I collagen and apatite crystals. The aim of this project is to gain a clearer understanding of the relationship between the organic and inorganic components of ivory dentine using analytical TEM, in order to utilise these analytical techniques in the context of common skeletal diseases such as osteoporosis and arthritis. TEM sections were prepared in both single and dual beam FIB instruments, using two standard lift-out techniques, in situ and ex situ. The FIB sections were systematically compared with sections prepared by ultramicrotomy, the traditional preparation route in biological systems, in terms of structural and chemical differences. A clear advantage of FIB milling over ultramicrotomy is that dehydration, embedding and section flotation can be eliminated, so that partial mineral loss due to dissolution is avoided. The characteristic banding of collagen fibrils was clearly seen in FIB milled sections without the need for any chemical staining, as is commonly employed in ultramicrotomy. The FIB milling technique was able to produce high-quality TEM sections of ivory dentine, which are suitable for further investigation using electron energy-loss spectroscopy (EELS) and energy-filtering TEM (EFTEM) to probe the collagen/apatite interface

Lozi, H., Goodwin, T.E., Rasmussen, L.E.L., Whitehouse, A.M., Schulte, B.A., 2009. Sexual dimorphism in the performance of chemosensory investigatory behaviours by African elephants (Loxodonta africana). Behaviour 146, 373-392.
Abstract: Sexual dimorphism in morphology can be accompanied by behavioural differences between the sexes. We examined if investigatory behaviour involving the trunk of African elephants showed sexual dimorphism. Males compete and search for females, but they have a lengthy period of development before they are socially viable mates. Receptive females are relatively rare. We hypothesized that males would display higher rates of chemosensory behaviour following puberty than females. Because males disperse, they were hypothesized to be more likely to contact elephants outside their kinship group. We observed the trunk tip, chemosensory behaviours of African elephants at Addo Elephant National Park, South Africa. For 208 elephants, we found no significant differences in state behaviours around waterholes by age or sex. Yet, older elephants were more likely to investigate the environment and elephant excrement than younger animals. Males were more likely to investigate urine and faeces than females. Only post-puberty animals contacted non-family with males investigating both sexes, while investigations by and to females only involved post-puberty males. Overall, the probability of performing chemosensory behaviours depended on age and sex. Male elephants appear more reliant than females on signals in urine and faeces with ensuing inspections of individuals through trunk tip contacts.

Manger, P.R., Pillay, P., Maseko, B.C., Bhagwandin, A., Gravett, N., Moon, D.J., Jillani, N., Hemingway, J., 2009. Acquisition of brains from the African elephant (Loxodonta africana): perfusion-fixation and dissection
113. J. Neurosci. Methods 179, 16-21.
Abstract: The current correspondence describes the in situ perfusion-fixation of the brain of the African elephant. Due to both the large size of proboscidean brains and the complex behaviour of these species, the acquisition of good quality material for comparative neuroanatomical analysis from these species is important. Three male African elephants (20-30 years) that were to be culled as part of a larger population management strategy were used. The animals were humanely euthanized and the head removed from the body. Large tubes were inserted into to the carotid arteries and the cranial vasculature flushed with a rapid (20 min) rinse of 100 l of cold saline (4 degrees C). Following the rinse the head was perfusion-fixed with a slower rinse (40 min) of 100 l of cold (4 degrees C) 4% paraformaldehyde in 0.1M phosphate buffer. This procedure resulted in well-fixed neural and other tissue. After perfusion the brains were removed from the skull with the aid of power tools, a procedure taking between 2 and 6h. The brains were immediately post-fixed in the same solution for 72 h at 4 degrees C. The brains were subsequently placed in a sucrose solution and finally an antifreeze solution and are stored in a -20 degrees C freezer. The acquisition of high quality neural material from African elephants that can be used for immunohistochemistry and electron microscopy is of importance in understanding the "hardware" underlying the behaviour of this species. This technique can be used on a variety of large mammals to obtain high quality material for comparative neuroanatomical studies

Murata, Y., Yonezawa, T., Kihara, I., Kashiwamura, T., Sugihara, Y., Nikaido, M., Okada, N., Endo, H., Hasegawa, M., 2009. Chronology of the extant African elephant species and case study of the species identification of the small African elephant with the molecular phylogenetic method
70. Gene 441, 176-186.
Abstract: Despite vigorous genetic studies of African elephants (Loxodonta africana and L. cyclotis) during the last decade, their evolutionary history is still obscure. Phylogenetic studies and coalescence time estimation using longer nucleotide sequence data from denser samplings are necessary to better understand the natural history of African elephants. Further, species identification among African elephants is sometimes very difficult using only the external morphological characteristics. This is a serious problem for making an adequate breeding plan in zoological gardens. In this paper, we investigated the continent-wide phylogeographical pattern of the African elephants and estimated the coalescence times among them. From these molecular data and geological evidence, we proposed an evolutionary scenario for the African elephants. We further demonstrated the effectiveness of molecular phylogenetic methods in species identification.

Paul, G., 2009. The nearly columnar limbs of elephants are very different from the more flexed, spring action limbs of running mammals and birds. J. Exp. Biol. 212, 152-3, author.

Saragusty, J., Hildebrandt, T.B., Behr, B., Knieriem, A., Kruse, J., Hermes, R., 2009. Successful cryopreservation of Asian elephant (Elephas maximus) spermatozoa. Anim Reprod. Sci. 115, 255-266.
Abstract: Reproduction in captive elephants is low and infant mortality is high, collectively leading to possible population extinction. Artificial insemination was developed a decade ago; however, it relies on fresh-chilled semen from just a handful of bulls with inconsistent sperm quality. Artificial insemination with frozen-thawed sperm has never been described, probably, in part, due to low semen quality after cryopreservation. The present study was designed with the aim of finding a reliable semen freezing protocol. Screening tests included freezing semen with varying concentrations of ethylene glycol, propylene glycol, trehalose, dimethyl sulfoxide and glycerol as cryoprotectants and assessing cushioned centrifugation, rapid chilling to suprazero temperatures, freezing extender osmolarity, egg yolk concentration, post-thaw dilution with cryoprotectant-free BC solution and the addition of 10% (v/v) of autologous seminal plasma. The resulting optimal freezing protocol uses cushioned centrifugation, two-step dilution with isothermal 285 m Osm/kg Berliner Cryomedium (BC) with final glycerol concentration of 7% and 16% egg yolk, and freezing in large volume by the directional freezing technique. After thawing, samples are diluted 1:1 with BC solution. Using this protocol, post-thaw evaluations results were: motility upon thawing: 57.2+/-5.4%, motility following 30 min incubation at 37 degrees C: 58.5+/-6.0% and following 3h incubation: 21.7+/-7.6%, intact acrosome: 57.1+/-5.2%, normal morphology: 52.0+/-5.8% and viability: 67.3+/-6.1%. With this protocol, good quality semen can be accumulated for future use in artificial inseminations when and where needed

Sherwood, C.C., Stimpson, C.D., Butti, C., Bonar, C.J., Newton, A.L., Allman, J.M., Hof, P.R., 2009. Neocortical neuron types in Xenarthra and Afrotheria: implications for brain evolution in mammals. Brain Struct. Funct. 213, 301-328.
Abstract: Interpreting the evolution of neuronal types in the cerebral cortex of mammals requires information from a diversity of species. However, there is currently a paucity of data from the Xenarthra and Afrotheria, two major phylogenetic groups that diverged close to the base of the eutherian mammal adaptive radiation. In this study, we used immunohistochemistry to examine the distribution and morphology of neocortical neurons stained for nonphosphorylated neurofilament protein, calbindin, calretinin, parvalbumin, and neuropeptide Y in three xenarthran species-the giant anteater (Myrmecophaga tridactyla), the lesser anteater (Tamandua tetradactyla), and the two-toed sloth (Choloepus didactylus)-and two afrotherian species-the rock hyrax (Procavia capensis) and the black and rufous giant elephant shrew (Rhynchocyon petersi). We also studied the distribution and morphology of astrocytes using glial fibrillary acidic protein as a marker. In all of these species, nonphosphorylated neurofilament protein-immunoreactive neurons predominated in layer V. These neurons exhibited diverse morphologies with regional variation. Specifically, high proportions of atypical neurofilament-enriched neuron classes were observed, including extraverted neurons, inverted pyramidal neurons, fusiform neurons, and other multipolar types. In addition, many projection neurons in layers II-III were found to contain calbindin. Among interneurons, parvalbumin- and calbindin-expressing cells were generally denser compared to calretinin-immunoreactive cells. We traced the evolution of certain cortical architectural traits using phylogenetic analysis. Based on our reconstruction of character evolution, we found that the living xenarthrans and afrotherians show many similarities to the stem eutherian mammal, whereas other eutherian lineages display a greater number of derived traits

Soltis, J., 2009. Vocal communication in African Elephants (Loxodonta africana)
61. Zoo. Biol. 28, 1-18.
Abstract: Research on vocal communication in African elephants has increased in recent years, both in the wild and in captivity, providing an opportunity to present a comprehensive review of research related to their vocal behavior. Current data indicate that the vocal repertoire consists of perhaps nine acoustically distinct call types, "rumbles" being the most common and acoustically variable. Large vocal production anatomy is responsible for the low-frequency nature of rumbles, with fundamental frequencies in the infrasonic range. Additionally, resonant frequencies of rumbles implicate the trunk in addition to the oral cavity in shaping the acoustic structure of rumbles. Long-distance communication is thought possible because low-frequency sounds propagate more faithfully than high-frequency sounds, and elephants respond to rumbles at distances of up to 2.5 km. Elephant ear anatomy appears designed for detecting low frequencies, and experiments demonstrate that elephants can detect infrasonic tones and discriminate small frequency differences. Two vocal communication functions in the African elephant now have reasonable empirical support. First, closely bonded but spatially separated females engage in rumble exchanges, or "contact calls," that function to coordinate movement or reunite animals. Second, both males and females produce "mate attraction" rumbles that may advertise reproductive states to the opposite sex. Additionally, there is evidence that the structural variation in rumbles reflects the individual identity, reproductive state, and emotional state of callers. Growth in knowledge about the communication system of the African elephant has occurred from a rich combination of research on wild elephants in national parks and captive elephants in zoological parks. Zoo Biol 28:1-18, 2009. (c) 2009 Wiley-Liss, Inc

Trimble, M.J., Ferreira, S.M., van Aarde, R.J., 2009. Drivers of megaherbivore demographic fluctuations: inference from elephants. Journal of Zoology 1-9.
Abstract: Environmentally induced variation in survival and fecundity generates demographic fluctuations that affect population growth rate. However, a general pattern of the comparative influence of variation in fecundity and juvenile survival on elephant population dynamics has not been investigated at a broad scale. We evaluated the relative importance of conception, gestation, first year survival and subsequent survivorship for controlling demographic variation by exploring the relationship between past environmental conditions determined by integrated normalized difference vegetation index (INDVI) and the shape of age distributions at 17 sites across Africa. We showed that, generally, INDVI during gestation best explained anomalies in age structure. However, in areas with low mean annual rainfall, INDVI during the first year of life was critical. The results challenge Eberhardt's paradigm for population analysis that suggests that populations respond to limited resource availability through a sequential decrease in juvenile survival, reproductive rate and adult survival. Contrastingly, elephants appear to respond first through a reduction in reproductive rate. We conclude that this discrepancy is likely due to the evolutionary significance of extremely large body size - an adaptation that increases survival rate but decreases reproductive potential. Other megaherbivores may respond similarly to resource limitation due to similarities in population dynamics. Knowing how vital rates vary with changing environmental conditions will permit better forecasts of the trajectories of megaherbivore populations.

Asher, R.J., Lehmann, T., 2008. Dental eruption in afrotherian mammals. BMC. Biol. 6, 14.
Abstract: BACKGROUND: Afrotheria comprises a newly recognized clade of mammals with strong molecular evidence for its monophyly. In contrast, morphological data uniting its diverse constituents, including elephants, sea cows, hyraxes, aardvarks, sengis, tenrecs and golden moles, have been difficult to identify. Here, we suggest relatively late eruption of the permanent dentition as a shared characteristic of afrotherian mammals. This characteristic and other features (such as vertebral anomalies and testicondy) recall the phenotype of a human genetic pathology (cleidocranial dysplasia), correlations with which have not been explored previously in the context of character evolution within the recently established phylogeny of living mammalian clades. RESULTS: Although data on the absolute timing of eruption in sengis, golden moles and tenrecs are still unknown, craniometric comparisons for ontogenetic series of these taxa show that considerable skull growth takes place prior to the complete eruption of the permanent cheek teeth. Specimens showing less than half (sengis, golden moles) or two-thirds (tenrecs, hyraxes) of their permanent cheek teeth reach or exceed the median jaw length of conspecifics with a complete dentition. With few exceptions, afrotherians are closer to median adult jaw length with fewer erupted, permanent cheek teeth than comparable stages of non-afrotherians. Manatees (but not dugongs), elephants and hyraxes with known age data show eruption of permanent teeth late in ontogeny relative to other mammals. While the occurrence of delayed eruption, vertebral anomalies and other potential afrotherian synapomorphies resemble some symptoms of a human genetic pathology, these characteristics do not appear to covary significantly among mammalian clades. CONCLUSION: Morphological characteristics shared by such physically disparate animals such as elephants and golden moles are not easy to recognize, but are now known to include late eruption of permanent teeth, in addition to vertebral anomalies, testicondy and other features. Awareness of their possible genetic correlates promises insight into the developmental basis of shared morphological features of afrotherians and other vertebrates

Carter, A.M., Miglino, M.A., Ambrosio, C.E., Santos, T.C., Rosas, F.C., Neto, J.A., Lazzarini, S.M., Carvalho, A.F., da Silva, V.M., 2008. Placentation in the Amazonian manatee (Trichechus inunguis). Reprod. Fertil. Dev. 20, 537-545.
Abstract: Evidence from several sources supports a close phylogenetic relationship between elephants and sirenians. To explore whether this was reflected in similar placentation, we examined eight delivered placentae from the Amazonian manatee using light microscopy and immunohistochemistry. In addition, the fetal placental circulation was described by scanning electron microscopy of vessel casts. The manatee placenta was zonary and endotheliochorial, like that of the elephant. The interhaemal barrier comprised maternal endothelium, cytotrophoblasts and fetal endothelium. We found columnar trophoblast beneath the chorionic plate and lining lacunae in this region, but there was no trace in the term placenta of haemophagous activity. The gross anatomy of the cord and fetal membranes was consistent with previous descriptions and included a four-chambered allantoic sac, as also found in the elephant and other afrotherians. Connective tissue septae descended from the chorionic plate and carried blood vessels to the labyrinth, where they gave rise to a dense capillary network. This appeared to drain into shorter vessels near the chorionic plate. The maternal vasculature could not be examined in the same detail, but maternal capillaries ran rather straight and roughly parallel to the fetal ones. Overall, there is a close resemblance in placentation between the manatee and the elephant

Drews, B., Hermes, R., Goritz, F., Gray, C., Kurz, J., Lueders, I., Hildebrandt, T.B., 2008. Early embryo development in the elephant assessed by serial ultrasound examinations. Theriogenology 69, 1120-1128.
Abstract: The elephant has an extraordinary long pregnancy, lasting 21 months. However, knowledge on embryo development is limited. To date, only single morphological observations of elephant embryo development associated with placentation are available, all lacking correlation to gestational age. The present study describes morphological characteristics of early embryo development in the elephant with exact biometric staging. Six pregnancies in five Asian and one African elephants with known conception dates were followed by 2D and 3D ultrasound, covering the embryonic period from ovulation to day 116 post-ovulation. The embryonic vesicle was earliest observed was on day 50 p.o. The proper embryo was not detected until day 62 p.o. Embryonic heartbeat was first observed on day 71 p.o. The allantois, which became visible as a single sacculation on day 71 p.o. was subdivided in four compartments on day 76 p.o. By day 95 p.o., head, rump, front and hind legs were clearly distinguished. Between days 95 and 103 p.o. the choriovitelline placenta was replaced by the chorioallantoic placenta. A physiological midgut herniation was transiently present between days 95 and 116 p.o. On the basis of the late appearance of the embryonic vesicle, delayed implantation in the elephant is discussed. The study provides a coherent description of elephant embryonic development, formation of the extraembryonic organs and their role in placenta formation, all of which are of interest for both comparative evolutionary studies and the improvement of assisted reproduction techniques

Hermes, R., Saragusty, J., Schaftenaar, W., Goritz, F., Schmitt, D.L., Hildebrandt, T.B., 2008. Obstetrics in elephants. Theriogenology 70, 131-144.
Abstract: Obstetrics, one of the oldest fields in veterinary medicine, is well described and practiced in domestic and exotic animals. However, when providing care during elephant birth or dystocia, veterinary intervention options differ greatly from any domestic species, and are far more limited due to the dimensions and specific anatomy of the elephant reproductive tract. In addition, aging of captive elephant populations and advanced age of primiparous females make active birth management increasingly important. Intrauterine infection, uterine inertia and urogenital tract pathologies are emerging as major causes for dystocia, often leading to foetal and dam death. This paper reviews the current knowledge on elephant birth and the factors associated with dystocia. It then summarises recommendations for birth and dystocia management. As Caesarean section, the most common ultima ratio in domestic animal obstetrics, is lethal and therefore not an option in the elephant, non-invasive medical treatment, induction of the Fergusson reflex or the conscious decision to leave a retained foetus until it is expelled voluntarily, are key elements in elephant obstetrics. Surgical strategies such as episiotomy and foetotomy are sometimes inevitable in order to try to save the life of the dam, however, these interventions result in chronic post-surgical complications or even fatal outcome. Limited reliable data on serum calcium concentrations, and pharmacokinetics and effect of exogenous oestrogen, oxytocin, and prostaglandins during birth provide the scope of future research, necessary to advance scientific knowledge on obstetrics in elephants

Liu, A.G., Seiffert, E.R., Simons, E.L., 2008. Stable isotope evidence for an amphibious phase in early proboscidean evolution. Proc. Natl. Acad. Sci. U. S. A 105, 5786-5791.
Abstract: The order Proboscidea includes extant elephants and their extinct relatives and is closely related to the aquatic sirenians (manatees and dugongs) and terrestrial hyracoids (hyraxes). Some analyses of embryological, morphological, and paleontological data suggest that proboscideans and sirenians shared an aquatic or semiaquatic common ancestor, but independent tests of this hypothesis have proven elusive. Here we test the hypothesis of an aquatic ancestry for advanced proboscideans by measuring delta(18)O in tooth enamel of two late Eocene proboscidean genera, Barytherium and Moeritherium, which are sister taxa of Oligocene-to-Recent proboscideans. The combination of low delta(18)O values and low delta(18)O standard deviations in Barytherium and Moeritherium matches the isotopic pattern seen in aquatic and semiaquatic mammals, and differs from that of terrestrial mammals. delta(13)C values of these early proboscideans suggest that both genera are likely to have consumed freshwater plants, although a component of C(3) terrestrial vegetation cannot be ruled out. The simplest explanation for the combined evidence from isotopes, dental functional morphology, and depositional environments is that Barytherium and Moeritherium were at least semiaquatic and lived in freshwater swamp or riverine environments, where they grazed on freshwater vegetation. These results lend new support to the hypothesis that Oligocene-to-Recent proboscideans are derived from amphibious ancestors

Miller, W., Drautz, D.I., Ratan, A., Pusey, B., Qi, J., Lesk, A.M., Tomsho, L.P., Packard, M.D., Zhao, F., Sher, A., Tikhonov, A., Raney, B., Patterson, N., Lindblad-Toh, K., Lander, E.S., Knight, J.R., Irzyk, G.P., Fredrikson, K.M., Harkins, T.T., Sheridan, S., Pringle, T., Schuster, S.C., 2008. Sequencing the nuclear genome of the extinct woolly mammoth. Nature 456, 387-390.
Abstract: In 1994, two independent groups extracted DNA from several Pleistocene epoch mammoths and noted differences among individual specimens. Subsequently, DNA sequences have been published for a number of extinct species. However, such ancient DNA is often fragmented and damaged, and studies to date have typically focused on short mitochondrial sequences, never yielding more than a fraction of a per cent of any nuclear genome. Here we describe 4.17 billion bases (Gb) of sequence from several mammoth specimens, 3.3 billion (80%) of which are from the woolly mammoth (Mammuthus primigenius) genome and thus comprise an extensive set of genome-wide sequence from an extinct species. Our data support earlier reports that elephantid genomes exceed 4 Gb. The estimated divergence rate between mammoth and African elephant is half of that between human and chimpanzee. The observed number of nucleotide differences between two particular mammoths was approximately one-eighth of that between one of them and the African elephant, corresponding to a separation between the mammoths of 1.5-2.0 Myr. The estimated probability that orthologous elephant and mammoth amino acids differ is 0.002, corresponding to about one residue per protein. Differences were discovered between mammoth and African elephant in amino-acid positions that are otherwise invariant over several billion years of combined mammalian evolution. This study shows that nuclear genome sequencing of extinct species can reveal population differences not evident from the fossil record, and perhaps even discover genetic factors that affect extinction

Organ, C.L., Schweitzer, M.H., Zheng, W., Freimark, L.M., Cantley, L.C., Asara, J.M., 2008. Molecular phylogenetics of mastodon and Tyrannosaurus rex. Science 320, 499.
Abstract: We report a molecular phylogeny for a nonavian dinosaur, extending our knowledge of trait evolution within nonavian dinosaurs into the macromolecular level of biological organization. Fragments of collagen alpha1(I) and alpha2(I) proteins extracted from fossil bones of Tyrannosaurus rex and Mammut americanum (mastodon) were analyzed with a variety of phylogenetic methods. Despite missing sequence data, the mastodon groups with elephant and the T. rex groups with birds, consistent with predictions based on genetic and morphological data for mastodon and on morphological data for T. rex. Our findings suggest that molecular data from long-extinct organisms may have the potential for resolving relationships at critical areas in the vertebrate evolutionary tree that have, so far, been phylogenetically intractable

Steenkamp, G., Ferguson, W.H., Boy, S.C., Ferreira, S.M., Bester, M.N., 2008. Estimating exposed pulp lengths of tusks in the African elephant (Loxodonta africana africana). J. S. Afr. Vet. Assoc. 79, 25-30.
Abstract: Captive and wild African elephants frequently suffer tusk fractures. Several institutions shorten the tusks of captive elephants to reduce fractures and injury as a result of behaviour within enclosures. Fracturing or coronal amputations that expose pulp lead to pain for the elephant. Estimating coronal pulp lengths may thus help to minimise the risk of pulp exposure during amputations. We aimed to determine the length of the pulp beyond the lip margin from an external tusk characteristic. Tusks collected from elephants in Namibia and the Kruger National Park had similar morphological relationships. This statistical property allowed us to correct for missing data in our data sets. Pulp volume and pulp length correlated with tusk circumference at the lip. Even so, the circumference at the lip could not predict the length of the pulp in the crown external to the lip. Our findings suggest that tusks, irrespective of sex or age, amputated further than 300 mm from the lip should not expose pulp

Thongtip, N., Saikhun, J., Mahasawangkul, S., Kornkaewrat, K., Pongsopavijitr, P., Songsasen, N., Pinyopummin, A., 2008. Potential factors affecting semen quality in the Asian elephant (Elephas maximus). Reprod. Biol. Endocrinol. 6, 9.
Abstract: BACKGROUND: One of the major obstacles in using artificial insemination to manage genetics of elephant population in captivity is the large variations in semen quality among ejaculates within the same and among individuals. The objectives of this study were to determine the influences of (1) age (2) seasonality (3) and circulating testosterone (SrTest), triiodothyronine (SrT3) and tetraiodothyronine (SrT4), as well as seminal (4) testosterone (SpTest), zinc (SpZn) and protein (SpTP) on semen quality in the Asian elephant METHODS: Analyses, including motility, viability and morphology were performed in semen samples collected twice monthly from 13 elephant bulls (age range, 10-to 72-years) by manual stimulation between July 2004 and June 2005. Serum samples obtained monthly were assessed for SrTest, SrT3, SrT4, and seminal plasma samples were evaluated for, SpTest, SpZn and SpTP. RESULTS: The highest semen quality was observed at age 23 to 43 years. Percentages of progressive motility and viable sperm were lowest at age 51 to 70 years (P < 0.05); on the other hand, sperm concentration was lowest at age 10 to 19 years (P < 0.05). Percentage of sperm with normal morphology was highest at age 23 to 43 years. The levels of SrT3, SrTest, SpTest and SpZn were lowest at age 51 to 70 years, whereas SrT4 was lowest at age 23 to 43 years. Seasonality significantly affected semen characteristics in which percentage of viable sperm and cell concentration were highest during rainy season and lowest during summer months (P < 0.05). However, percentage of sperm with normal morphology was highest in summer and lowest in rainy season (P < 0.05). Seasonality significantly influenced SrTest with elevated concentrations observed in rainy season and winter (P < 0.05). CONCLUSION: This study indicates that age and seasonality had influence on semen characteristics in the Asian elephant. The knowledge obtained in this study will improve our understanding of the reproductive biology of this species

Valeix, M., Fritz, H., Matsika, R., Matsvimbo, F., Madzikanda, H., 2008. The role of water abundance, thermoregulation, perceived predation risk and interference competition in water access by African herbivores. African Journal of Ecology 46, 402-410.
Abstract: In African savannas, surface water can become limiting and an understanding of how animals address the trade-offs between different constraints to access this resource is needed. Here, we describe water access by ten African herbivore species in Hwange National Park, Zimbabwe. and we explore four possible determinants of the observed behaviours: water abundance, thermoregulation, perceived predation risk and interference competition. On average, herbivores were observed to drink in 80% of visits to a waterhole. The probability of drinking was higher in 2003 (474 mm) than in 2004 (770 mm), and at the end of the dry season than at its beginning. For larger species, this probability may also be related to risks of interference competition with elephants or other herbivores. For smaller species, this probability may also be related to the perceived risk of predation. We also investigate the time spent accessing water to drink. The influence of herd size and the presence of young on the time spent accessing water for most species suggests that perceived predation risk plays a role. Themoregulation also affects this time: during the hottest periods, herbivores spend less time in open areas. unless when wind is strong, probably owing to evapotranspired heat loss.

Bates, L.A., Byrne, R.W., 2007. Creative or created: using anecdotes to investigate animal cognition. Methods 42, 12-21.
Abstract: In non-human animals, creative behaviour occurs spontaneously only at low frequencies, so is typically missed by standardised observational methods. Experimental approaches have tended to rely overly on paradigms from child development or adult human cognition, which may be inappropriate for species that inhabit very different perceptual worlds and possess quite different motor capacities than humans. The analysis of anecdotes offers a solution to this impasse, provided certain conditions are met. To be reliable, anecdotes must be recorded immediately after observation, and only the records of scientists experienced with the species and the individuals concerned should be used. Even then, interpretation of a single record is always ambiguous, and analysis is feasible only when collation of multiple records shows that a behaviour pattern occurs repeatedly under similar circumstances. This approach has been used successfully to study a number of creative capacities of animals: the distribution, nature and neural correlates of deception across the primate order; the occurrence of teaching in animals; and the neural correlates of several aptitudes--in birds, foraging innovation, and in primates, innovation, social learning and tool-use. Drawing on these approaches, we describe the use of this method to investigate a new problem, the cognition of the African elephant, a species whose sheer size and evolutionary distance from humans renders the conventional methods of comparative psychology of little use. The aim is both to chart the creative cognitive capacities of this species, and to devise appropriate experimental methods to confirm and extend previous findings

Bouley, D.M., Alarcón, C.N., Hildebrandt, T., O'connell-Rodwell, C.E., ., 2007. The distribution, density and three-dimensional histomorphology of Pacinian corpuscles in the foot of the Asian elephant (Elephas maximus) and their potential role in seismic communication. J Anat 211, 428-435.
Abstract: Both Asian (Elephas maximus) and African (Loxodonta africana) elephants produce low-frequency, high-amplitude rumbles that travel well through the ground as seismic waves, and field studies have shown that elephants may utilize these seismic signals as one form of communication. Unique elephant postures observed in field studies suggest that the elephants use their feet to 'listen' to these seismic signals, but the exact sensory mechanisms used by the elephant have never been characterized. The distribution, morphology and tissue density of Pacinian corpuscles, specialized mechanoreceptors, were studied in a forefoot and hindfoot of Asian elephants. Pacinian corpuscles were located in the dermis and distal digital cushion and were most densely localized to the anterior, posterior, medial and lateral region of each foot, with the highest numbers in the anterior region of the forefoot (52.19%) and the posterior region of the hindfoot (47.09%). Pacinian corpuscles were encapsulated, had a typical lamellar structure and were most often observed in large clusters. Three-dimensional reconstruction through serial sections of the dermis revealed that individual Pacinian corpuscles may be part of a cluster. By studying the distribution and density of these mechanoreceptors, we propose that Pacinian corpuscles are one possible anatomic mechanism used by elephants to detect seismic waves.

Gunga, H.C., Suthau, T., Bellmann, A., Friedrich, A., Schwanebeck, T., Stoinski, S., Trippel, T., Kirsch, K., Hellwich, O., 2007. Body mass estimations for Plateosaurus engelhardti using laser scanning and 3D reconstruction methods. Naturwissenschaften 94, 623-630.
Abstract: Both body mass and surface area are factors determining the essence of any living organism. This should also hold true for an extinct organism such as a dinosaur. The present report discusses the use of a new 3D laser scanner method to establish body masses and surface areas of an Asian elephant (Zoological Museum of Copenhagen, Denmark) and of Plateosaurus engelhardti, a prosauropod from the Upper Triassic, exhibited at the Paleontological Museum in Tubingen (Germany). This method was used to study the effect that slight changes in body shape had on body mass for P. engelhardti. It was established that body volumes varied between 0.79 m(3) (slim version) and 1.14 m(3) (robust version), resulting in a presumable body mass of 630 and 912 kg, respectively. The total body surface areas ranged between 8.8 and 10.2 m(2), of which, in both reconstructions of P. engelhardti, approximately 33% account for the thorax area alone. The main difference between the two models is in the tail and hind limb reconstruction. The tail of the slim version has a surface area of 1.98 m(2), whereas that of the robust version has a surface area of 2.73 m(2). The body volumes calculated for the slim version were as follows: head 0.006 m(3), neck 0.016 m(3), fore limbs 0.020 m(3), hind limbs 0.08 m(3), thoracic cavity 0.533 m(3), and tail 0.136 m(3). For the robust model, the following volumes were established: 0.01 m(3) head, neck 0.026 m(3), fore limbs 0.025 m(3), hind limbs 0.18 m(3), thoracic cavity 0.616 m(3), and finally, tail 0.28 m(3). Based on these body volumes, scaling equations were used to assess the size that the organs of this extinct dinosaur have

Hildebrandt, T., Drews, B., Gaeth, A.P., Goeritz, F., Hermes, R., Schmitt, D., Gray, C., Rich, P., Streich, W.J., Short, R.V., Renfree, M.B., 2007. Foetal age determination and development in elephants. Proc. Biol. Sci. 274, 323-331.
Abstract: Elephants have the longest pregnancy of all mammals, with an average gestation of around 660 days, so their embryonic and foetal development have always been of special interest. Hitherto, it has only been possible to estimate foetal ages from theoretical calculations based on foetal mass. The recent development of sophisticated ultrasound procedures for elephants has now made it possible to monitor the growth and development of foetuses of known gestational age conceived in captivity from natural matings or artificial insemination. We have studied the early stages of pregnancy in 10 captive Asian and 9 African elephants by transrectal ultrasound. Measurements of foetal crown-rump lengths have provided the first accurate growth curves, which differ significantly from the previous theoretical estimates based on the cube root of foetal mass. We have used these to age 22 African elephant foetuses collected during culling operations. Pregnancy can be first recognized ultrasonographically by day 50, the presumptive yolk sac by about day 75 and the zonary placenta by about day 85. The trunk is first recognizable by days 85-90 and is distinct by day 104, while the first heartbeats are evident from around day 80. By combining ultrasonography and morphology, we have been able to produce the first reliable criteria for estimating gestational age and ontological development of Asian and African elephant foetuses during the first third of gestation.

Kinahan, A.A., Pimma, S.L., van Aarde, R.J., 2007. Ambient temperature as a determinant of landscape use in the savanna elephant, Loxodonta africana. Journal of Thermal Biology 32, 47-58.
Abstract: Elephants occur in landscapes where temperatures can reach 50 degrees C. Due to their large size they may face physiological problems of dissipating heat during such high temperatures. In spite of this, no one seems to have considered ambient temperature as limiting landscape choices in elephants. We recorded hourly landscape use in free-ranging elephants using GPS collars. We also placed temperature data loggers in each of the landscapes, to obtain corresponding ambient temperatures for each hour. Our results suggest that elephants may select landscapes based on the rate at which temperatures changed and also for shade. We suggest that these selected variables provide a thermal benefit to individuals. As such, we propose that landscape use in elephants may be constrained by their thermal physiological requirements as well as other resources such as food and water.

Murphy, W.J., Pringle, T.H., Crider, T.A., Springer, M.S., Miller, W., 2007. Using genomic data to unravel the root of the placental mammal phylogeny. Genome Res. 17, 413-421.
Abstract: The phylogeny of placental mammals is a critical framework for choosing future genome sequencing targets and for resolving the ancestral mammalian genome at the nucleotide level. Despite considerable recent progress defining superordinal relationships, several branches remain poorly resolved, including the root of the placental tree. Here we analyzed the genome sequence assemblies of human, armadillo, elephant, and opossum to identify informative coding indels that would serve as rare genomic changes to infer early events in placental mammal phylogeny. We also expanded our species sampling by including sequence data from >30 ongoing genome projects, followed by PCR and sequencing validation of each indel in additional taxa. Our data provide support for a sister-group relationship between Afrotheria and Xenarthra (the Atlantogenata hypothesis), which is in turn the sister-taxon to Boreoeutheria. We failed to recover any indels in support of a basal position for Xenarthra (Epitheria), which is suggested by morphology and a recent retroposon analysis, or a hypothesis with Afrotheria basal (Exafricoplacentalia), which is favored by phylogenetic analysis of large nuclear gene data sets. In addition, we identified two retroposon insertions that also support Atlantogenata and none for the alternative hypotheses. A revised molecular timescale based on these phylogenetic inferences suggests Afrotheria and Xenarthra diverged from other placental mammals approximately 103 (95-114) million years ago. We discuss the impacts of this topology on earlier phylogenetic reconstructions and repeat-based inferences of phylogeny

O'Connell-Rodwell, C.E., 2007. Keeping an "ear" to the ground: seismic communication in elephants. Physiology (Bethesda) 287-294.
Abstract: This review explores the mechanisms that elephants may use to send and receive seismic signals from a physical, anatomical, behavioral, and physiological perspective. The implications of the use of the vibration sense as a multimodal signal will be discussed in light of the elephant's overall fitness and survival.

Roca, A.L., Georgiadis, N., O'Brien, S.J., 2007. Cyto-nuclear genomic dissociation and the African elephant species question. Quat. Int. 169-170, 4-16.
Abstract: Studies of skull morphology and of nuclear DNA have strongly concluded that African elephants comprise two species. Nonetheless, Debruyne (2005) has suggested a single-species model for Loxodonta based on the polyphyly of a single genetic locus, mitochondrial DNA (mtDNA). Discordant patterns between mitochondrial and nuclear DNA markers were subsequently reported in some African savanna elephant populations, further supporting a two-species model, and prompting us to re-examine here the geographic distribution of different elephant morphotypes and their relationship to nuclear and mtDNA phylogeographic patterns. We used exact tests to compare the distribution of forest elephant-typical and savanna elephant-typical characteristics across eight published datasets containing morphological, mtDNA or nuclear DNA data for African elephants. Among the elephants examined by Debruyne (2005), we found that patterns of forest vs. savanna characteristics were significantly different (p < 10(-5)) between mtDNA and morphology, suggesting the presence of cyto-nuclear genomic dissociation. We show that the eight African elephant continent-wide datasets compared, including that of Debruyne (2005), together support a two-species model with cyto-nuclear genomic dissociation rather than a one-species model, and together indicate that Africa harbors two species of elephant

Tabuce, R., Marivaux, L., Adaci, M., Bensalah, M., Hartenberger, J.L., Mahboubi, M., Mebrouk, F., Tafforeau, P., Jaeger, J.J., 2007. Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade. Proc. Biol. Sci. 274, 1159-1166.
Abstract: The phylogenetic pattern and timing of the radiation of mammals, especially the geographical origins of major crown clades, are areas of controversy among molecular biologists, morphologists and palaeontologists. Molecular phylogeneticists have identified an Afrotheria clade, which includes several taxa as different as tenrecs (Tenrecidae), golden moles (Chrysochloridae), elephant-shrews (Macroscelididae), aardvarks (Tubulidentata) and paenungulates (elephants, sea cows and hyracoids). Molecular data also suggest a Cretaceous African origin for Afrotheria within Placentalia followed by a long period of endemic evolution on the Afro-Arabian continent after the mid-Cretaceous Gondwanan breakup (approx. 105-25 Myr ago). However, there was no morphological support for such a natural grouping so far. Here, we report new dental and postcranial evidence of Eocene stem hyrax and macroscelidid from North Africa that, for the first time, provides a congruent phylogenetic view with the molecular Afrotheria clade. These new fossils imply, however, substantial changes regarding the historical biogeography of afrotheres. Their long period of isolation in Africa, as assumed by molecular inferences, is now to be reconsidered inasmuch as Eocene paenungulates and elephant-shrews are here found to be related to some Early Tertiary Euramerican 'hyopsodontid condylarths' (archaic hoofed mammals). As a result, stem members of afrotherian clades are not strictly African but also include some Early Paleogene Holarctic mammals

Witter, K., Egger, G.F., Boeck, P., 2007. Renaut bodies in nerves of the trunk of the African elephant, Loxodonta africana. J. Morphol. 268, 414-422.
Abstract: Renaut bodies are loosely textured, cell-sparse structures in the subperineurial space of peripheral nerves, frequently found at sites of nerve entrapment. The trunk of the elephant is a mobile, richly innervated organ, which serves for food gathering, object grasping and as a tactile organ. These functions of the trunk lead to distortion and mechanical compression of its nerves, which can therefore be expected to contain numerous Renaut bodies. Samples of the trunk wall of an adult African elephant (Loxodonta africana) were examined histologically using conventional staining methods, immunohistochemistry, and lectin histochemistry. Architecture of nerve plexuses and occurrence of Renaut bodies in the elephant trunk were compared with those in tissues surrounding the nasal vestibule of the pig. Prominent nerve plexuses were found in all layers of the elephant trunk. Almost all (81%) nerve profiles contained Renaut bodies, a basophilic, discrete subperineurial layer resembling cushions around the nerve core. In contrast, Renaut bodies were seen in only 15% of nerve profiles in the porcine nasal vestibule. Within Renaut bodies, fusiform fibroblasts and round, ruff-like cells were placed into a matrix of acidic glycosaminoglycans with delicate collagen and very few reticular fibers. The turgor of this matrix is thought to protect nerves against compression and shearing strain. Renaut bodies are readily stained with alcian blue (pH 2.5) favorably in combination with immunohistochemical markers of nerve fibers. They should be regarded as a physiological response to repeated mechanical insults and are distinct from pathological alterations. alterations

Allen, W.R., 2006. Ovulation, pregnancy, placentation and husbandry in the African elephant (Loxodonta africana)
470. Philos. Trans. R. Soc. Lond B Biol. Sci. 361, 821-834.
Abstract: The African elephant reproduces so efficiently in the wild that overpopulation is now a serious problem in some game parks in Zimbabwe, Botswana and South Africa. The female reaches puberty between 10 and 12 years of age in the wild and, when in captivity, shows oestrous cycles of 14-15 weeks duration. She readily conceives a singleton in the wild yet her uterus has the capacity for twins. She shows a gestation length of 22 months and, in the wild, shows a population density and feed dependent intercalving interval of 4-8 years. The trophoblast erodes the lumenal epithelium of the endometrium and stimulates upgrowths of blood vessel-containing stromal villi, which develop eventually into the broad, tightly folded lamellae of the zonary, endotheliochorial placenta. Significant quantities of leaked maternal erythrocytes and ferric iron are phagocytosed by specialized trophoblast cells in the haemophagous zones at the lateral edges of the placental band. Although the placenta itself is endocrinologically inert, the foetal gonads, which enlarge greatly during the second half of pregnancy can synthesize 5alpha-dihydryoprogesterone and other 5alpha pregnane derivatives from cholesterol and pregnenolone. These products may synergize with progestagens secreted by the 2-8 large corpora lutea which are always present in the maternal ovaries throughout gestation to maintain the pregnancy state

Benz, A., Zenker, W., Hildebrandt, T.B., Weissengruber, G., Geyer, H. Recent findings about the macroscopic and microscopic morphology of the elephants hooves (Elephantidae). Proceedings International Elephant Conservation & Research Symposium. 38-41. 2006. 2006.
Ref Type: Conference Proceeding

Bertschinger, H., Delsink, A., Kirkpatrick, J.F., Human, A., Grobler, D., van Altena, J.J. Management of elephant populations in private South African game reserves with porcine zona pellucida vaccine. 2006 Proceedings American Association of Zoo Veterinarians. 283-285. 2006.
Ref Type: Conference Proceeding
Abstract: Control of African elephant populations has become an absolute necessity in a number of game reserves in southern Africa. The two main methods used to control populations so far are culling and translocation. Culling, besides being regarded as inhumane and unacceptable in many quarters, is not suitable for smaller populations. It requires that whole family units are culled simultaneously which could mean that in reserves with 10 to 50 elephants a considerable portion, if not the entire population, is killed. As far as translocation is concerned, limited new space is available for elephants. The only alternative to the two above options is to control the rate of reproduction. The porcine zona pellucida (pZP) vaccine has been used to successfully contracept wild horses and other wildlife species. Work on the contraception of African elephants was initiated in the Kruger National Park in 1995 when the potential for using the porcine zona pellucida (pZP) was investigated. Subsequently the first field trials on wild elephants were carried out in Kruger and the results clearly showed that elephants could be contracepted with the pZP vaccine, although the efficacy achieved was 80%. During these field trials safety and reversibility werecould be demonstrated. In 2000 an elephant contraceptive program was initiated at Makalali Private Game Reserve, RSA, which has become the flagship model for immunocontrol in African elephants. The preliminary findings have been reported in three publications.During the first year, all 18 cows that were individually identified and older than 12 yr of age were treated. During the next 4 yr the number of cows contracepted increased to 23 as young animals were added to the program. The standard vaccination procedure during the first year consisted of a primary vaccination (600 μg or 400 μg pZP with 0.5 ml Freund's modified complete adjuvant) followed by boosters (200 μg pZP with 0.5 ml Freund's incomplete adjuvant) at 3 to 6-wk intervals. Annual boosters to maintain antibody titers and contraceptive effect followed. To date, the success rate on cows that have passed reserve-specific intercalving period of 56 mo has been 100%. The population stabilized within 3 yr by which time when all cows that had been pregnant at the time of first vaccination in 2000 had calved. Once again safety during pregnancy (14 cows pregnant at 2-21 mo gestation when first treated gave birth to normal healthy calves) as well as side effects that were limited to occasional lumps at the site of vaccination could be shown. Following ground darting, behavioral patterns returned to pre-darting status within 2 days. During 2003 and 2004 most boosters were administered from a helicopter; whereas, previously they had been done from a vehicle or on foot. In all cases, drop-out darts were used. Time taken for vaccination from helicopter take-off to landing was about 30 min (1.5 min per cow; 30 min for total time). This required prior knowledge of the locations of family units or that an individual in each unit is radio-collared. Herds settled down much more quickly (1-2 days) than if darted from the ground. Since then we have vaccinated another 107 elephant cows in eight game reserves. The cow populations have ranged from 4 to 43. In one of the reserves, Mabula, RSA, two of the four cows vaccinated have passed the mean intercalving intervals of the reserve with neither of them producing a calf. Treatment at the remaining reserves was initiated in 2004 or 2005 and it is too early to evaluate results. The most difficult reserve in terms of the vaccination process was Welgevonden, RSA, (35 000 ha) with 43 cows. The reserve is mountainous and heavily wooded. None of the elephants were collared and individuals could not be easily identified on the day of primary vaccination. The total flying time during which individuals were identified and vaccinated was 4.5 hr. Administration of the first booster took about 2 hr to locate and vaccinate each cow. Between the first and second booster the first rains occurred, followed by the spring flush of the vegetation. By the time the second booster was attempted late in November, the trees all had foliage. Only half the cows were located and darted because the elephants were very difficult to spot under the tree canopies. The valuable lessons we learned from this were: 1) that helicopter vaccinations should be performed when most trees are bare, and 2) when larger populations are vaccinated repeatedly during the first year, one cow in each family unit should be radio-collared. This makes rapid location of each unit possible and cuts down on the major cost factor that is flying time. Elephant behavior is being monitored in all eight reserves where contraception is being applied. Because most of them have been contracepted recently, only the data from Makalali is available. The elephants at Makalali have been monitored intensively almost on a daily basis. To date, no anomalies in terms of aggressive or indifferent behavior with regards to nursing time, nursing behavior and calf proximity have been noted. No change in the cows' social hierarchy has been noted. Since January 2003, a total of 15 heats were observed in 10 cows (nine in 2003 and six in 2004) with four mating episodes. For the same period, 38 musth occasions were seen in five bulls (26 in 2003 and 12 in 2004). These occasions include musth displayed in the same bull during consecutive days or within the same musth cycle. The greatest occurrence of musth was recorded in the largest, dominant bull. Bulls were not observed harassing or separating cows off from their herds or calves as a result of increased estrous frequency. Thus, the Makalali program demonstrates that pZP does not cause herd fragmentation, harassment by bulls, change in rank and other negative behaviors normally associated with hormonal contraceptives. In conclusion we feel that it is important to emphasize the following points: The pZP vaccine can be used successfully to contracept African elephants The vaccine is safe during pregnancy and has no negative effect on birth or calf raising It has no side effects other than occasional swelling at the site of vaccination It is reversible Other than an increased incidence of heat no behavioral side effects were seen.

Drews, B., Göritz, F., Hermes, R., Streich, J.W., Rich, P., Schmitt, D., Lung, N., Renfree, M.B., Gaeth, A.P., Short, R.V., Hildebrandt, T.B. Morphological and ultrasonographic characterization of the embryonic development in elephants. Proceedings International Elephant Conservation & Research Symposium. 82-83. 2006. 2006.
Ref Type: Conference Proceeding

Fischer, M.S., Blickhan, R., 2006. The tri-segmented limbs of therian mammals: kinematics, dynamics, and self-stabilization--a review
397. J. Exp. Zool. A Comp Exp. Biol. 305, 935-952.
Abstract: The evolution of therian mammals is to a large degree marked by changes in their motion systems. One of the decisive transitions has been from the sprawled, bi-segmented to the parasagittal, tri-segmented limb. Here, we review aspects of the tri-segmented limb in locomotion which have been elucidated in our research groups in the last 10 years. First, we report the kinematics of the tri-segmented therian limb from mouse to elephant in order to explore general principles of the therian limb configuration and locomotion. Torques will be reported from a previous paper (Witte et al., 2002. J Exp Biol 205:1339-1353) for a better understanding of the anti-gravity work of all limb joints. The stability of a limb in z-configuration will be explained and its advantage with respect to other potential solutions from modeling will be discussed. Finally, we describe how the emerging concept of self-stability can be explained for a tri-segmented leg template and how it affects the design of the musculoskeletal system and the operation of legs during locomotion. While locomotion has been considered as mainly a control problem in various disciplines, we stress the necessity to reduce control as much as possible. Central control can be cheap if the limbs are "intelligent" by means of their design. Gravity-induced movements and self-stability seem to be energy-saving mechanisms

Hofreiter, M., Lister, A., 2006. Mammoths
459. Curr. Biol. 16, R347-R348.

Schmitt, D., Cartmill, M., Griffin, T.M., Hanna, J.B., Lemelin, P., 2006. Adaptive value of ambling gaits in primates and other mammals
460. J. Exp. Biol. 209, 2042-2049.
Abstract: At speeds between the walk and the gallop, most mammals trot. Primates almost never trot, and it has been claimed that they transition directly from a walk to a gallop without any distinctive mid-speed running gait. If true, this would be another characteristic difference between the locomotion of primates and that of most other quadrupedal mammals. Presently, however, few data exist concerning the actual presence or absence of intermediate-speed gaits (i.e. gaits that are used between a walk and a gallop) in primates. Video records of running in twelve primate species reveal that, unlike most other mammals, all the primates studied almost exclusively adopt an 'amble'--an intermediate-speed running gait with no whole-body aerial phase--rather than trot. Ambling is also common in elephants and some horses, raising the question of why ambling is preferred over trotting in these diverse groups of animals. Mathematical analyses presented here show that ambling ensures continuous contact of the body with the substrate while dramatically reducing vertical oscillations of the center of mass. This may explain why ambling appears to be preferable to trotting for extremely large terrestrial mammals such as elephants and for arboreal mammals like primates that move on unstable branches. These findings allow us to better understand the mechanics of these unusual running gaits and shed new light on primate locomotor evolution

Shoshani, J., Kupsky, W.J., Marchant, G.H., 2006. Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution. Brain Res Bull 70, 124-157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.

Shoshani, J., Walter, R.C., Abraha, M., Berhe, S., Tassy, P., Sanders, W.J., Marchant, G.H., Libsekal, Y., Ghirmai, T., Zinner, D., 2006. A proboscidean from the late Oligocene of Eritrea, a "missing link" between early Elephantiformes and Elephantimorpha, and biogeographic implications
384. Proc. Natl. Acad. Sci. U. S. A 103, 17296-17301.
Abstract: We report on a late Oligocene proboscidean species from Eritrea, dated to 26.8 +/- 1.5 Mya. This "missing link" between early elephantiformes and Elephantimorpha is the oldest known nongomphothere proboscidean to probably display horizontal tooth displacement, typical of elephants [Elephantimorpha consists of Mammutida (mastodons) and Elephantida, and Elephantida includes gomphotheres, stegodons, and elephants]. Together with the newly discovered late Oligocene gomphotheres from Chilga, Ethiopia, the Eritrean taxon points to the importance of East Africa as a major area for the knowledge of the early evolution of Elephantimorpha before the faunal exchange between Eurasia and Africa

Shoshani, J., Kupsky, W.J., Marchant, G.H., 2006. Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution
446. Brain Res. Bull. 70, 124-157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior

Siegal-Willott, J., Isaza, R., Johnson, R., Blaik, M. Clinical evaluation of distal limb radiography and growth plate closure in the juvenile Asian elephant (Elephas maximus).
2006 Proceedings American Association of Zoo Veterinarians. 181-182. 2006.
Ref Type: Conference Proceeding
Abstract: The thoracic limb digits of 11 healthy juvenile Asian elephants (Elephas maximus) were evaluated radiographically to assess normal developmental anatomy. Parameters evaluated included: the location(s) of centers of ossification, relative age at time of phalangeal ossification, and relative times of growth plate closure in the bones of the distal forelimb. Specifically, the third phalanx (P3) of each digit was evaluated, as well as the first (P1) and second (P2) phalanges of the third digit (D3). A retrospective evaluation of radiographs from juvenile elephants was also done to augment the data set. This study reports the methods used to obtain high-quality radiographs of the elephant foot, the locations of centers of ossification based on radiographic evaluation, and the relative times of growth plate closure within the digital bones. The settings used to obtain the radiographs used in this study for P3 are presented in Table 1. Radiographs of D3, P1, and P2 were obtained in a similar manner, using a 45° angle for focal spot positioning. The kilovoltage power and milliampere seconds were adjusted as needed. Radiographic evaluation of the juvenile Asian elephants revealed variability in the shape of P3 based on age of the animal and degree of ossification of P3. The relative times of growth plate closure and number of ossifications were also determined. The information presented will help clinicians in radiographing elephants, interpreting foot radiographs, and recognizing normal versus abnormal anatomy. It will also help in aging juvenile elephants, investigating diseases and deaths, and recognizing normal patterns of toe and foot development.

Takahashi, H., Yamashita, M., Shigehara, N., 2006. Cranial photographs of mammals on the web: The Mammalian Crania Photographic Archive (MCPA2) and a comparison of bone image databases. Anthropological Science 114, 217-222.
Abstract: The Mammalian Crania Photographic Archive (MCPA2) is a website (http://1kai.dokkyomed.ac.jp/mammal/en/mammal.html) that includes a collection of 10,950 photographs of mammalian crania, which have been taken with a high-resolution digital camera. In the present report, we outline the characteristics of MCPA2 and how it was created, and make brief comparisons with several similar websites currently accessible via the internet. The archived MCPA2 materials include 1825 cranial specimens, ranging from insectivores to elephants, which have been macerated in Japan during the past 35 years and prepared for osteological study. Of the 16 orders represented in the database, primates comprise the major group with 704 specimens. Each cranium was placed with the orbitomeatal (Frankfort) or palatine plane horizontal, and was photographed in six perpendicular views from a long distance using a telephoto or telemacro lens. These long-distance shots decrease perspective distortion that lead to measurement errors when studying cranial profiles and landmark positions, and enable detailed observation and measurement of specific bony characteristics on a computer screen. From our website, images can be searched using (1) the taxonomic table, (2) Japanese name, (3) English name, and (4) scientific name. In the page of search results, in addition to the images, four caliper measurements and additional text (taxonomy, sex, and age) are available for every specimen.

Uni, S., Bain, O., Agatsuma, T., Katsumi, A., Baba, M., Yanai, T., Takaoka, H., 2006. New filarial nematode from Japanese serows (Naemorhedus crispus: Bovidae) close to parasites from elephants
403. Parasite 13, 193-200.
Abstract: A new onchocercid species, Loxodontofilaria caprini n. sp. (Filarioidea: Nematoda), found in subcutaneous tissues of 37 (33%) of 112 serows (Noemorhedus crispus) examined in Japan, is described. The female worm had the characteristics of Loxodontofilaria, e.g., the large body size, well-developed esophagus with a shallow buccal cavity, and the long tail with three caudal lappets. The male worm of the new species, which was first described in the genus, had unequal length of spicules, 10 pairs of pre- and post-caudal papillae, and three terminal caudal lappets. Deirids were present in both sexes. Among four species of the genus loxodontofiloria: one from the hippopotamus and three from the Elepantidae, L. caprini n. sp. appears close to L. asiatica Bain, Baker & Chabaud, 1982, a subcutaneous parasite of Elephas indicus in Myanmar (Burma). However, L. caprini n. sp. is distinct from L. asiatica in that the Japanese female worm has an esophagus half as long and the microfilariae also half as long with a coiled posterior. The microfilariae were found in the skin of serows. The new parasite appears to clearly illustrate a major event in the evolution of onchocercids: the host-switching. This might have occurred on the Eurasian continent, where elephantids and the lineage of rupicaprines diversified during the Pliocene-Pleistocene, or in Japan, into which some of these hosts migrated

Weissengruber, G.E., Fuss, F.K., Egger, G., Stanek, G., Hittmair, K.M., Forstenpointner, G., 2006. The elephant knee joint: morphological and biomechanical considerations
513. Journal of Anatomy 208, 59-72.
Abstract: Elephant limbs display unique morphological features which are related mainly to supporting the enormous body weight of the animal. In elephants, the knee joint plays important roles in weight bearing and locomotion, but anatomical data are sparse and lacking in functional analyses. In addition, the knee joint is affected frequently by arthrosis. Here we examined structures of the knee joint by means of standard anatomical techniques in eight African (Loxodonta africana) and three Asian elephants (Elephas maximus). Furthermore, we performed radiography in five African and two Asian elephants and magnetic resonance imaging (MRI) in one African elephant. Macerated bones of 11 individuals (four African, seven Asian elephants) were measured with a pair of callipers to give standardized measurements of the articular parts. In one Asian and three African elephants, kinematic and functional analyses were carried out using a digitizer and according to the helical axis concept. Some peculiarities of healthy and arthrotic knee joints of elephants were compared with human knees. In contrast to those of other quadruped mammals, the knee joint of elephants displays an extended resting position. The femorotibial joint of elephants shows a high grade of congruency and the menisci are extremely narrow and thin. The four-bar mechanism of the cruciate ligaments exists also in the elephant. The main motion of the knee joint is extension-flexion with a range of motion of 142 degrees . In elephants, arthrotic alterations of the knee joint can lead to injury or loss of the cranial (anterior) cruciate ligament

Weissengruber, G.E., Egger, G.F., Hutchinson, J.R., Groenewald, H.B., Elsasser, L., Famini, D., Forstenpointner, G., 2006. The structure of the cushions in the feet of African elephants (Loxodonta africana)
380. Journal of Anatomy 209, 781-792.
Abstract: The uniquely designed limbs of the African elephant, Loxodonta africana, support the weight of the largest terrestrial animal. Besides other morphological peculiarities, the feet are equipped with large subcutaneous cushions which play an important role in distributing forces during weight bearing and in storing or absorbing mechanical forces. Although the cushions have been discussed in the literature and captive elephants, in particular, are frequently affected by foot disorders, precise morphological data are sparse. The cushions in the feet of African elephants were examined by means of standard anatomical and histological techniques, computed tomography (CT) and magnetic resonance imaging (MRI). In both the forelimb and the hindlimb a 6th ray, the prepollex or prehallux, is present. These cartilaginous rods support the metacarpal or metatarsal compartment of the cushions. None of the rays touches the ground directly. The cushions consist of sheets or strands of fibrous connective tissue forming larger metacarpal/metatarsal and digital compartments and smaller chambers which were filled with adipose tissue. The compartments are situated between tarsal, metatarsal, metacarpal bones, proximal phalanges or other structures of the locomotor apparatus covering the bones palmarly/plantarly and the thick sole skin. Within the cushions, collagen, reticulin and elastic fibres are found. In the main parts, vascular supply is good and numerous nerves course within the entire cushion. Vater-Pacinian corpuscles are embedded within the collagenous tissue of the cushions and within the dermis. Meissner corpuscles are found in the dermal papillae of the foot skin. The micromorphology of elephant feet cushions resembles that of digital cushions in cattle or of the foot pads in humans but not that of digital cushions in horses. Besides their important mechanical properties, foot cushions in elephants seem to be very sensitive structures

Zuba, J.R., Oosterhuis, J.E., Pessier, A.P. The toenail "abscess" in elephants: treatment options including cryotherapy and pathologic similarities with equine proliferative pododermatitis (canker). 2006 Proceedings American Association of Zoo Veterinarians. 187-190. 2006.
Ref Type: Conference Proceeding
Abstract: Foot problems potentially represent the single most important clinical disease of captive elephants. Predisposing factors include obesity, lack of exercise, nail or sole overgrowth, improper foot care, poor hygiene, inappropriate enclosure surfaces, poor conformation, malnutrition and secondary skeletal disorders such as degenerative joint disease. Furthermore, factors such as elephant management philosophy, disposition of elephants, facilities and competency of staff in caring for elephant feet will contribute significantly to the foot health of captive animals. It is important to note that these conditions are rarely reported in free-ranging elephants. The elephant toenail abscess is characterized grossly by proliferative outgrowth of "crab meat-like" tissue that may acutely rupture through the surface of the nail wall and/or adjacent cuticle or sole. True abscess formation with localized collections of suppurative material is not a consistent clinical feature. In most cases, the inciting cause of these lesions are typically not found and are likely due to one or more of the predisposing factors listed above. Once established, these frustrating lesions require extensive, intensive and prolonged medical attention. If not cared for properly, these wounds may progress to phalangeal osteomyelitis and the need for surgical intervention. Sole abscesses are equally frustrating and difficult to manage with proposed etiologies similar to toenail lesions. There are no reports in the literature describing the pathology of the classic proliferative abscess tissue of the elephant nail abscess. Although variously interpreted as fibrous or granulation tissue, the authors are unaware of previous histologic descriptions of this tissue. Biopsy samples of toenail abscess tissue from two Asian elephants (Elephas maximus) at the San Diego Wild Animal Park (SDWAP) consisted of stratified squamous epithelium arranged in columns resembling horn tubules. The predominant histologic finding was marked, near diffuse, hydropic degeneration of keratinocytes. There were multifocal areas of suppurative inflammation with admixed bacterial colonies. Inflammatory foci comprised only a small portion of the lesion and were interpreted as the external surfaces of the biopsy with likely secondary bacterial colonization. Because descriptions of the normal histology of the elephant toenail could not be located, a grossly normal toenail from a different Asian elephant was obtained to compare histologic features with those of the toenail abscesses. Sections demonstrated formation of the toenail in a manner similar to that of the hoof of the horse and cattle with tubular, intertubular and laminar horn. Primary and secondary epidermal laminae were identified. Proliferative lesions of horn-producing epithelium associated with ballooning degeneration and inadequate keratinization of keratinocytes, have been described in horses as equine "canker" and coronary band dystrophy. Equine canker is most commonly observed in the hind feet of draft horses and begins in the frog sometimes with extension to the sole and hoof wall. Grossly, lesions are characterized by soft white papillary to "cauliflower-like" tissue associated with a foul odor. Similar to what is noted in elephant foot problems, predisposing factors for the development of equine canker include poor hygiene or wet environmental conditions. There is a lack of gross and histologic description of the normal nail and sole tissue of the elephant and further investigations are warranted. A review of the anatomy and histology of the normal equine hoof may provide a basic understanding of the elephant nail until more specific and detailed elephant information is available. From our investigation, the authors offer that a more accurate description of the elephant toenail abscess would be proliferative pododermatitis, the term synonymous with equine canker. A more colloquial term such as "elephant canker" may be appropriate, as well. Canker in the horse is an uncommon but difficult to treat disease of the hoof. Historically, treatment options for elephant toenail abscesses include corrective trimming, superficial debridement and application of topical disinfectants or antibiotics. Others have constructed innovative sandals to treat and protect the affected sole or nail with success. The use of regional intravenous perfusion of the affected limb with antibiotics has also been successful. Since the elephant nail abscess now appears to be histologically and clinically comparable to equine canker, this novel characterization of an old disease may offer unique insight for treatment. In the least, it has provided our practice with a new list of treatment options and experienced equine clinicians for consultation who have been managing patients with a similar disease for many years. One of the Asian elephants at the SDWAP has had chronic toenail abscesses for over 2 yr. Radiographs of the affected digits, as reported by others to assess degree of involvement, have fortunately been negative for evidence of osteomyelitis. Several bacterial and fungal cultures of deep tissue biopsies and swabs of affected lesions have resulted in a mixture of organisms with no consistent single etiologic agent. Biopsies were found negative for presence of viral DNA (elephant papillomavirus and herpesvirus) by PCR. Typical elephant foot care at the SDWAP includes trimming and debriding with hoof knives, foot soaks and topical antibiotics. Although difficult, attempts are made in keeping the affected foot clean and dry. Following recommendations for the treatment of equine canker, we recently implemented the routine use of cryotherapy in all elephants with proliferative pododermatitis with improved success in the control and recession of exuberant nail lesions. The proliferative tissue of the nail is first cleaned then disinfected, debrided, trimmed with hoof knives and allowed to dry. Modified brass branding tools with contact surfaces of variable size (2-5 cm diameter) and shape (round or ovoid) are placed into liquid nitrogen (-196 C) for several minutes and then placed directly on the cankerous tissue for 30-60 sec. This process is then repeated 4-5 min later, following a complete thaw of tissue. Within 24 hr, the cryoburned tissue becomes macerated and necrotic and is readily removed with gentle scrubbing. Cryotherapy offers the advantage of destroying tissue to a deeper level than trimming alone and provides hemostasis, as well. Because of decreased sensation at the cryotherapy treatment site, a memorable painful event is avoided and the elephant patient is more routinely accepting of this technique. With the use of hoof knives, we typically remove 2-3 mm of proliferative tissue before the patient refuses further treatment, presumably due to discomfort. With cryotherapy, we are able to remove an additional 3-5 mm of tissue by cell freezing and necrosis. The result is quicker resolution of cankerous lesions without the need for aggressive, and potentially painful, interventions. In conclusion, it appears that elephant nail abscesses can best be described as proliferative pododermatitis, or canker, as is seen in other species. Further gross and microscopic descriptions of normal and pathologic nail or sole lesions are necessary. Routine cryotherapy has shown promise in the treatment of these chronic, frustrating and potentially devastating lesions of our captive elephants.

Benz, A., Zenker, W., Hildebrandt, T.B., Weissengruber, G.E., Geyer, H. About the macroscopic and microscopic morphology of elephants' hooves (Elephantidae). Erkrankungen der Zootiere. Verhandlungsbericht des Internationalen Symposiums über die Erkrankungen der Zoo- und Wildtiere / International Symposium on Diseases of Zoo and Wild Animals. 42, 164-166. 2005.
Ref Type: Conference Proceeding

Benz, A. The elephant's hoof: macroscopic and microscopic morphology of defined locations under consideration of pathological changes. 2005. Vetsuisse-Fakultät Universität Zürich.
Ref Type: Thesis/Dissertation

Edwards, H.G., Jorge Villar, S.E., Nik Hassan, N.F., Arya, N., O'Connor, S., Charlton, D.M., 2005. Ancient biodeterioration: an FT-Raman spectroscopic study of mammoth and elephant ivory
576. Anal. Bioanal. Chem. 383, 713-720.
Abstract: Raman spectra of mammoth ivory specimens have been recorded using near-infrared excitation, and comparisons made with modern Asian and African elephant ivories. Whereas the most ancient mammoth ivory (60-65 ky) showed no evidence for an organic collagen component, more recent samples of mammoth ivory indicated that some preservation had occurred, although with biodeterioration of the protein structure exhibited by the amide I and III bands in the 1200-1700 cm(-1) region of the Raman spectrum. The consequent difficulties encountered when applying chemometrics methods to ancient ivory analysis (which are successful for modern specimens) are noted. In the most ancient mammoth ivory specimens, which are extensively fragmented, evidence of mineralization is seen, with the production of gypsum, calcite and limonite; Raman microscopic analysis of crystalline material inside the fissures of the mammoth ivory shows the presence of gypsum as well as cyanobacterial colonisation. The application of Raman spectroscopy to the nondestructive analysis of archaeological materials in order to gain information of relevance to their preservation or restoration is highlighted

Glickman, S.E., Short, R.V., Renfree, M.B., 2005. Sexual differentiation in three unconventional mammals: spotted hyenas, elephants and tammar wallabies
566. Horm. Behav. 48, 403-417.
Abstract: The present review explores sexual differentiation in three non-conventional species: the spotted hyena, the elephant and the tammar wallaby, selected because of the natural challenges they present for contemporary understanding of sexual differentiation. According to the prevailing view of mammalian sexual differentiation, originally proposed by Alfred Jost, secretion of androgen and anti-Mullerian hormone (AMH) by the fetal testes during critical stages of development accounts for the full range of sexually dimorphic urogenital traits observed at birth. Jost's concept was subsequently expanded to encompass sexual differentiation of the brain and behavior. Although the central focus of this review involves urogenital development, we assume that the novel mechanisms described in this article have potentially significant implications for sexual differentiation of brain and behavior, a transposition with precedent in the history of this field. Contrary to the "specific" requirements of Jost's formulation, female spotted hyenas and elephants initially develop male-type external genitalia prior to gonadal differentiation. In addition, the administration of anti-androgens to pregnant female spotted hyenas does not prevent the formation of a scrotum, pseudoscrotum, penis or penile clitoris in the offspring of treated females, although it is not yet clear whether the creation of masculine genitalia involves other steroids or whether there is a genetic mechanism bypassing a hormonal mediator. Wallabies, where sexual differentiation occurs in the pouch after birth, provide the most conclusive evidence for direct genetic control of sexual dimorphism, with the scrotum developing only in males and the pouch and mammary glands only in females, before differentiation of the gonads. The development of the pouch and mammary gland in females and the scrotum in males is controlled by genes on the X chromosome. In keeping with the "expanded" version of Jost's formulation, secretion of androgens by the fetal testes provides the best current account of a broad array of sex differences in reproductive morphology and endocrinology of the spotted hyena, and androgens are essential for development of the prostate and penis of the wallaby. But the essential circulating androgen in the male wallaby is 5alpha androstanediol, locally converted in target tissues to DHT, while in the pregnant female hyena, androstenedione, secreted by the maternal ovary, is converted by the placenta to testosterone (and estradiol) and transferred to the developing fetus. Testicular testosterone certainly seems to be responsible for the behavioral phenomenon of musth in male elephants. Both spotted hyenas and elephants display matrilineal social organization, and, in both species, female genital morphology requires feminine cooperation for successful copulation. We conclude that not all aspects of sexual differentiation have been delegated to testicular hormones in these mammals. In addition, we suggest that research on urogenital development in these non-traditional species directs attention to processes that may well be operating during the sexual differentiation of morphology and behavior in more common laboratory mammals, albeit in less dramatic fashion

Hakeem, A.Y., Hof, P.R., Sherwood, C.C., Switzer, R.C., III, Rasmussen, L.E., Allman, J.M., 2005. Brain of the African elephant (Loxodonta africana): neuroanatomy from magnetic resonance images. Anat. Rec. A Discov. Mol. Cell Evol. Biol. 287, 1117-1127.
Abstract: We acquired magnetic resonance images of the brain of an adult African elephant, Loxodonta africana, in the axial and parasagittal planes and produced anatomically labeled images. We quantified the volume of the whole brain (3,886.7 cm3) and of the neocortical and cerebellar gray and white matter. The white matter-to-gray matter ratio in the elephant neocortex and cerebellum is in keeping with that expected for a brain of this size. The ratio of neocortical gray matter volume to corpus callosum cross-sectional area is similar in the elephant and human brains (108 and 93.7, respectively), emphasizing the difference between terrestrial mammals and cetaceans, which have a very small corpus callosum relative to the volume of neocortical gray matter (ratio of 181-287 in our sample). Finally, the elephant has an unusually large and convoluted hippocampus compared to primates and especially to cetaceans. This may be related to the extremely long social and chemical memory of elephants

Konishi, S., 2005. [Jaws of herbivorous mammals]
582. Clin. Calcium 15, 1414-1417.
Abstract: The jaws of herbivorous mammals are characterized by their large occlusal surface of the molar; high crown of the molar; long snout; etc. However, elephants, the biggest herbivorous mammal, have other characteristics. In the evolutionary trends of proboscidean skulls, concomitant with the increase in tusk size comes on the enlargement, antero-posterior shortening, dorso-ventral elongation of the cranium with increasing cheek teeth size. Naturally, the jaw follows the same evolutionary trends as the cranium

Luikart, K.A., Stover, S.M., 2005. Chronic sole ulcerations associated with degenerative bone disease in two Asian elephants (Elephas maximus). J. Zoo. Wildl. Med. 36, 684-688.
Abstract: Chronic foot lesions and degenerative joint disease are common causes of morbidity in elephants. Lesions may become intractable and progressive despite intensive treatment regimens. The forelimbs of two Asian elephants (Elephas maximus) with chronic nonhealing sole ulcerations were examined using manual dissection and computed tomography. Both elephants had abnormal limb conformation that preceded the development of sole ulcerations. In both cases, sole ulcers were associated with remodeling and degeneration of underlying bones of the digits. Conformational abnormalities and altered weight distribution in these individuals may have induced compensatory bony degeneration and secondary ulcer formation. Sole ulcerations associated with digital abnormalities may have a guarded prognosis for resolution, even with aggressive treatment. Because limb conformational abnormalities could predispose to or result from chronic digital lesions, elephants with conformational abnormalities may have increased likelihood of having chronic sole ulcerations

Mobasheri, A., Gent, T.C., Womack, M.D., Carter, S.D., Clegg, P.D., Barrett-Jolley, R., 2005. Quantitative analysis of voltage-gated potassium currents from primary equine (Equus caballus) and elephant (Loxodonta africana) articular chondrocytes
618. Am. J. Physiol Regul. Integr. Comp Physiol 289, R172-R180.
Abstract: In this comparative study, we have established in vitro models of equine and elephant articular chondrocytes, examined their basic morphology, and characterized the biophysical properties of their primary voltage-gated potassium channel (Kv) currents. Using whole cell patch-clamp electrophysiological recording from first-expansion and first-passage cells, we measured a maximum Kv conductance of 0.15 +/- 0.04 pS/pF (n = 10) in equine chondrocytes, whereas that in elephant chondrocytes was significantly larger (0.8 +/- 0.4 pS/pF, n = 4, P </= 0.05). Steady-state activation parameters of elephant chondrocytes (V = -22 +/- 6 mV, k = 11.8 +/- 3 mV, n = 4) were not significantly different from those of horse chondrocytes (V = -12.5 +/- 4.3 mV, k = 12 +/- 2, n = 10). This suggests that there would be slightly more resting Kv activation in elephant chondrocytes than in their equine counterparts. Kinetic analysis revealed that both horse and elephant chondrocyte Kv currents had similar activation and inactivation parameters. Pharmacological investigation of equine chondrocyte Kv currents showed them to be powerfully inhibited by the potassium channel blockers tetraethylammonium and 4-aminopyridine but not by dendrotoxin-I. Immunohistochemical studies using polyclonal antibodies to Kv1.1-Kv1.5 provided evidence for expression of Kv1.4 in equine chondrocytes. This is the first electrophysiological study of equine or elephant chondrocytes. The data support the notion that voltage-gated potassium channels play an important role in regulating the membrane potential of articular chondrocytes and will prove useful in future modeling of electromechanotransduction of fully differentiated articular chondrocytes in these and other species

Natiello, M., Lewis, P., Samuelson, D., 2005. Comparative anatomy of the ciliary body of the West Indian manatee (Trichechus manatus) and selected species. Vet. Ophthalmol. 8, 375-385.
Abstract: OBJECTIVE: To examine the anatomy of the ciliary body in the West Indian manatee (Trichechus manatus), paying close attention to its vascularization and to compare to those of its distant relative, the African elephant (Loxodonta africana), the amphibious hippopotamus (Hippopotamus amphibius) and the aquatic short-finned pilot whale (Globicephala macrorhynchus). PROCEDURE: Specimens from each species were preserved in 10% buffered formalin, and observed stereomicroscopically before being embedded in paraffin, sectioned and stained by Masson trichrome, hematoxylin and eosin, and periodic acid-Schiff for light microscopic evaluation. RESULTS: The network of blood vessels in the ciliary processes of the West Indian manatee appear to have an intricate pattern, especially with regard to venous outflow. Those of the elephant are slightly less complex, while those of the hippopotamus and whale have different vascular patterns within the ciliary body. Musculature within the ciliary body is absent within the manatee and pilot whale. CONCLUSIONS: In general, there appears to be a direct relationship between the increased development of vasculature and the loss of musculature within the ciliary bodies of the aquatic and amphibious mammals presently studied. Specifically, the ciliary body of the West Indian manatee has a comparatively unique construction, especially with regard to its vasculature.

Perez-Barberia, F.J., Gordon, I.J., 2005. Gregariousness increases brain size in ungulates
586. Oecologia. 145, 41-52.
Abstract: The brain's main function is to organise the physiological and behavioural responses to environmental and social challenges in order to keep the organism alive. Here, we studied the effects that gregariousness (as a measurement of sociality), dietary habits, gestation length and sex have on brain size of extant ungulates. The analysis controlled for the effects of phylogeny and for random variability implicit in the data set. We tested the following groups of hypotheses: (1) Social brain hypothesis-gregarious species are more likely to have larger brains than non-gregarious species because the former are subjected to demanding and complex social interactions; (2) Ecological hypothesis-dietary habits impose challenging cognitive tasks associated with finding and manipulating food (foraging strategy); (3) Developmental hypotheses (a) energy strategy: selection for larger brains operates, primarily, on maternal metabolic turnover (i.e. gestation length) in relation to food quality because the majority of the brain's growth takes place in utero, and finally (b) sex hypothesis: females are expected to have larger brains than males, relative to body size, because of the differential growth rates of the soma and brain between the sexes. We found that, after adjusting for body mass, gregariousness and gestation length explained most of the variation in brain mass across the ungulate species studied. Larger species had larger brains; gregarious species and those with longer gestation lengths, relative to body mass, had larger brains than non-gregarious species and those with shorter gestation lengths. The effect of diet was negligible and subrogated by gestation length, and sex had no significant effect on brain size. The ultimate cause that could have triggered the co-evolution between gestation length and brain size remains unclear

Pichardo, M., 2005. Taxonomic revision of Central Mexican mammoths in Paleoindian sites
522. Anthropol. Anz. 63, 409-413.
Abstract: Central Mexican mammoth species taxonomy has been based on the quotient Molar length/Number of dental plates, which sorted three species, Mammuthus imperator, columbi and ?jeffersonii. New evidence from skull morphology sorts only two subspecies, M. columbi columbi and M. columbi felicis as being present during Paleoindian time

Raubenheimer, E.J., Ngwenya, S.P., 2005. The role of ivory in the survival of the African elephant
510. SADJ. 60, 426, 430.
Abstract: The unique chequered pattern of polished ivory has created a perverted commercial demand for elephant tusks. The morphologic basis of the pattern, which makes ivory a sought after product for the manufacturing of works of art, is discussed. Chemical analyses of ivory holds great potential in tracing the source of illegally harvested tusks and exposing poorly managed elephant sanctuaries. The impact of uncontrolled ivory hunting on the population genetics of the African elephant is briefly reviewed

Roth, G., Dicke, U., 2005. Evolution of the brain and intelligence
612. Trends Cogn Sci. 9, 250-257.
Abstract: Intelligence has evolved many times independently among vertebrates. Primates, elephants and cetaceans are assumed to be more intelligent than 'lower' mammals, the great apes and humans more than monkeys, and humans more than the great apes. Brain properties assumed to be relevant for intelligence are the (absolute or relative) size of the brain, cortex, prefrontal cortex and degree of encephalization. However, factors that correlate better with intelligence are the number of cortical neurons and conduction velocity, as the basis for information-processing capacity. Humans have more cortical neurons than other mammals, although only marginally more than whales and elephants. The outstanding intelligence of humans appears to result from a combination and enhancement of properties found in non-human primates, such as theory of mind, imitation and language, rather than from 'unique' properties

Shoshani, J., Tassy, P., 2005. Advances in proboscidean taxonomy & classification, anatomy & physiology, and ecology & behavior. Quaternary International 126-128, 5-20.
Abstract: With the addition of 13 new taxa, we recognized 175 species and subspecies of proboscideans, classified in 42 genera and 10 families. The three extant species are: forest African elephant (Loxodonta cyclotis), bush African elephant (L. africana), and Asian elephant (Elephas maximus, with three subspecies). Rigorous analysis of characters published or awaiting publication is imperative for better understanding of the cladistic relationships among currently recognized proboscideans. Here we focus on ''aquatic ancestry'' of Proboscidea, interordinal relationships within Placentalia, proboscidean taxonomy in general and South American in particular, anatomy and physiology and some ecological considerations. New taxa above the family level include sister taxa Mammutida and Elephantida, and Plesielephantiformes as a sister taxon to Elephantiformes. Neontological research is currently under way on the hyoid apparatus, lungs, brain, hearing, ecology and behavior. Topics for future research include: phylogenetic positions of anthracobunids, Moeritherium, tetralophodont gomphotheres, Stegolophodon and Stegodon, and intra-familial relationships among Loxodonta, Elephas and Mammuthus, and continuing studies on encephalization quotient. Certain anatomical features and functions (e.g., the hyoid apparatus that helps in food procurement, in production of infrasonic sounds, and in storing water to be used in time of stress) evolved about 25 million years ago, in time for diversification into new niches when grasses appeared in the landscape.

Surovell, T., Waguespack, N., Brantingham, P.J., 2005. Global archaeological evidence for proboscidean overkill
616. Proc. Natl. Acad. Sci. U. S. A 102, 6231-6236.
Abstract: One million years ago, proboscideans occupied most of Africa, Europe, Asia, and the Americas. Today, wild elephants are only found in portions of sub-Saharan Africa and South Asia. Although the causes of global Pleistocene extinctions in the order Proboscidea remain unresolved, the most common explanations involve climatic change and/or human hunting. In this report, we test the overkill and climate-change hypotheses by using global archaeological spatiotemporal patterning in proboscidean kill/scavenge sites. Spanning approximately 1.8 million years, the archaeological record of human subsistence exploitation of proboscideans is preferentially located on the edges of the human geographic range. This finding is commensurate with global overkill, suggesting that prehistoric human range expansion resulted in localized extinction events. In the present and the past, proboscideans have survived in refugia that are largely inaccessible to human populations

Weissengruber, G.E., Egerbacher, M., Forstenpointner, G., 2005. Structure and innervation of the tusk pulp in the African elephant (Loxodonta africana)
617. Journal of Anatomy 206, 387-393.
Abstract: African elephants (Loxodonta africana) use their tusks for digging, carrying and behavioural display. Their healing ability following traumatic injury is enormous. Pain experience caused by dentin or pulp damage of tusks seems to be negligible in elephants. In this study we examined the pulp tissue and the nerve distribution using histology, electron microscopy and immunhistochemistry. The results demonstrate that the pulp comprises two differently structured regions. Randomly orientated collagen fibres characterize a cone-like part lying rostral to the foramen apicis dentis. Numerous nerve fibres and Ruffini endings are found within this cone. Rostral to the cone, delicate collagen fibres and large vessels are orientated longitudinally. The rostral two-thirds of the pulp are highly vascularized, whereas nerve fibres are sparse. Vessel and nerve fibre distribution and the structure of connective tissue possibly play important roles in healing and in the obviously limited pain experience after tusk injuries and pulp alteration. The presence of Ruffini endings is most likely related to the use of tusks as tools

Zack, S.P., Penkrot, T.A., Bloch, J.I., Rose, K.D., 2005. Affinities of 'hyopsodontids' to elephant shrews and a Holarctic origin of Afrotheria
622. Nature 434, 497-501.
Abstract: Macroscelideans (elephant shrews or sengis) are small-bodied (25-540 g), cursorial (running) and saltatorial (jumping), insectivorous and omnivorous placental mammals represented by at least 15 extant African species classified in four genera. Macroscelidea is one of several morphologically diverse but predominantly African placental orders classified in the superorder Afrotheria by molecular phylogeneticists. The distribution of modern afrotheres, in combination with a basal position for Afrotheria within Placentalia and molecular divergence-time estimates, has been used to link placental diversification with the mid-Cretaceous separation of South America and Africa. Morphological phylogenetic analyses do not support Afrotheria and the fossil record favours a northern origin of Placentalia. Here we describe fossil postcrania that provide evidence for a close relationship between North American Palaeocene-Eocene apheliscine 'hyopsodontid' 'condylarths' (early ungulates or hoofed mammals) and extant Macroscelidea. Apheliscine postcranial morphology is consistent with a relationship to other ungulate-like afrotheres (Hyracoidea, Proboscidea) but does not provide support for a monophyletic Afrotheria. As the oldest record of an afrothere clade, identification of macroscelidean relatives in the North American Palaeocene argues against an African origin for Afrotheria, weakening support for linking placental diversification to the break-up of Gondwana

Boellaard, J.W., Schlote, W., Hofer, W., 2004. Species-specific ultrastructure of neuronal lipofuscin in hippocampus and neocortex of subhuman mammals and humans
630. Ultrastruct. Pathol. 28, 341-351.
Abstract: Lipofuscin represents an integral part of neurons and glial cells in mammals and in submammalian species. It is a special lysosomal organelle, takes part of cellular metabolism, and is a structural expression of catabolic pathways. Species-specific differences of lipofuscin indicate metabolic differences of the relevant neurons. The authors have studied the ultrastructure of neuronal lipofuscin in the hippocampus and cerebral neocortex of dogs, horses, cows, elephants, rats, mice, apes, and humans to answer the question of species-specific differences of this organelle. Paraffin sections of formalin-fixed material were investigated by hematoxylin-eosin and PAS staining, by fluorescence microscopy for autofluorescence, with a laser scanning confocal microscope and by electron microscopy. In the animals studied and in humans the lipofuscin displayed, in addition to the general trilaminar substructure, species-specific appearances. No differences were found in the lipofuscin structure between neocortical and hippocampal neurons of the separate animal species. In contrast, in humans, neurons of the hippocampus showed a particular lipofuscin structure, not only different from the neocortical one, but also with differences between CA1 and CA3/4 sectors. Interestingly, in apes a transitional situation was found with slight differences between neocortical and hippocampal lipofuscin, especially in the rhesus monkey. This peculiarity was corroborated by the distribution of special pentilaminar linear structures in the lipofuscin pigment in all animals, only sparsely in the rhesus monkey and not in humans. The results indicate that lipofuscin ultrastructure of neocortical and hippocampal neurons is species specific and that lipofuscin in the human hippocampal neurons displays structures characteristic of man differing from the neocortical neuronal lipofuscin. The neuronal lipofuscin of apes, especially of the rhesus monkey displays structures in between humans and lower mammals. Nothing is known about the functional significance of these findings. They may indicate metabolic and/or functional characteristics of the relevant neurons

Boy, S.C., Steenkamp, G., 2004. Neural innervation of the tusk pulp of the African elephant (Loxodonta africana)
730. Veterinary Record 154, 372-374.

Brown, J.L., Walker, S.L., Moeller, T., 2004. Comparative endocrinology of cycling and non-cycling Asian (Elephas maximus) and African (Loxodonta africana) elephants
729. Gen. Comp Endocrinol. 136, 360-370.
Abstract: Up to 14% of Asian and 29% of African elephants in captivity are not cycling normally or exhibit irregular cycles based on progestin profiles. To determine if ovarian acyclicity is related to other disruptions in endocrine activity, serum pituitary, thyroid, adrenal, and ovarian hormones in weekly samples collected for 6-25 months were compared between normal cycling (n=22 each species) and non-cycling (n=6 Asian; n=30 African) elephants. A subset of cycling females (n=4 Asian, 7 African) also were blood sampled daily during the follicular phase to characterize the peri-ovulatory period. In normal cycling females, two leutinizing hormone (LH) surges were observed 3 weeks apart during a normal follicular phase, with the second inducing ovulation (ovLH). Serum FSH concentrations were highest at the beginning of the non-luteal phase, declining to nadir concentrations within 4 days of the ovLH surge. FSH remained low until after the ovLH surge and then increased during the luteal phase. A species difference was noted in prolactin secretion. In the African elephant, prolactin was increased during the follicular phase, but in Asian elephants concentrations remained stable throughout the cycle. Patterns of thyroid hormones (thyroid-stimulating hormone, TSH; free and total thyroxine, T4; free and total triiodothyronine, T3) and cortisol secretion were not affected by estrous cycle stage or season in cycling elephants. In non-cycling elephants, there were no fluctuating patterns of LH, FSH, or prolactin secretion. Overall mean concentrations of all hormones were similar to those in cycling animals, with the exception of FSH, prolactin, and estradiol. Mean serum FSH concentrations were lower due to females not exhibiting normal cyclic increases, whereas serum estradiol was higher overall in most acyclic females. Prolactin concentrations were significantly increased in 11 of 30 non-cycling females, all of which were African elephants. In sum, while there were no consistent endocrine anomalies associated with ovarian acyclicity, hyperprolactinemia may be one cause of ovarian dysfunction. The finding of elevated estrogens in some acyclic females also deserves further investigation, especially determining how it relates to reproductive tract pathologies

Carter, A.M., Enders, A.C., Kunzle, H., Oduor-Okelo, D., Vogel, P., 2004. Placentation in species of phylogenetic importance: the Afrotheria
705. Anim Reprod. Sci. 82-83, 35-48.
Abstract: Afrotheria, one of four mammalian superorders, comprises elephants, sea cows, hyraxes, aardvark, elephant shrews, tenrecs and golden moles. Their placentas either form an equatorial band or are discoid in shape. The interhemal region, separating fetal and maternal blood, is endotheliochorial in elephants, aardvark and possibly the sea cows, but hemochorial in the remaining orders. There is a secondary epitheliochorial placenta in elephant shrews while a similar structure in tenrecs erodes maternal tissues. Specialized hemophagous regions are a striking characteristic of some of these placentas yet absent in hyraxes, elephant shrews, and golden moles. It is possible that the common ancestor of the Afrotheria had an endotheliochorial placenta. Establishment of a hemochorial condition, as seen in rock hyraxes, elephant shrews, tenrecs, and golden moles, would be a more recent development. The elephant, manatee, and aardvark all have circumferential placentas. Thus the formation of a discoid placenta with a more or less extensive secondary placenta in elephant shrews and tenrecs would also be a derived state

Debruyne, R., 2004. [Contribution of molecular phylogeny and morphometrics to the systematics of African elephants]
590. J. Soc. Biol. 198, 335-342.
Abstract: African elephants are conventionally classified as a single species: Loxodonta africana (Blumenbach 1797). However, the discovery in 1900 of a smaller form of the African elephant, spread throughout the equatorial belt of this land, has given rise to a debate over the relevance of a second species of elephant in Africa. The twentieth century has not provided any definite answer to this question. Actually, recent molecular analyses have sustained this issue by advocating either a division of forest elephants into a valid species, or their inclusion as a subspecies of L. africana. Our work initiated at the National Museum of Natural History of Paris provides new molecular (mitochondrial) and morphological (and morphometrical) evidence making it possible to propose a comprehensive phylogenetic hypothesis. It appears that there is no conclusive argument to keep forest elephants (cyclotis form) and savannah elephants (africana form) apart in two distinct species. A high level of mitochondrial introgression between the two forms, as well as a continuum in the morphology of the skulls of the two morphotypes rather suggests that, despite an ancient division, these two taxa freely interbreed wherever their ranges intersect. We thus adopt a conservative systematic position in considering these two forms as two subspecies, respectively: L. africana africana, the savannah elephant, and L. africana cyclotis, the forest elephant. We finally discuss the conservation topic in the light of this systematic framework

Enders, A.C., Carter, A.M., 2004. What can comparative studies of placental structure tell us?--A review
739. Placenta 25 Suppl A , S3-S9.
Abstract: The diversity of placental structures in Eutherian mammals is such that drawing generalizations from the definitive forms is problematic. There are always areas of reduced interhaemal distance whether the placenta is epitheliochorial, synepitheliochorial, endotheliochorial or haemochorial. However, the thinning may be achieved by different means. The presence of a haemophagous area as an iron transport facilitator is generally associated with endotheliochorial placentae but is also found in sheep and goats (synepitheliochorial) and in tenrecs and hyaenas (haemochorial). Although similar chorioallantoic placentae are found within families, structure begins to diverge at the ordinal level and there is little correlation at the supraordinal level of phylogeny. Differences in formation and function of the yolk sac provide additional variation. There would appear to be considerable adaptive pressure for development or retention of the haemochorial type of chorioallantoic placenta. This type of placenta has several possible drawbacks including more ready passage of fetal cells to the maternal organism and, should the haemochorial condition be achieved early, oxidative stress. At any rate no animal larger than the human and gorilla has this type of placenta. The endotheliochorial condition is found in animals as large as the bears, manatee and elephants. In addition to the ungulates, the epitheliochorial condition is present in the largest animals with the longest gestation periods, the whales. Considering the length of time since the early stages of mammalian evolution, it is probable that few unmodified structural features are present in any currently surviving mammal. Nevertheless, more complete studies of divergent types of mammalian placenta should help our understanding of mammalian interrelationships as well as placental function

Gobbel, L., Fischer, M.S., Smith, T.D., Wible, J.R., Bhatnagar, K.P., 2004. The vomeronasal organ and associated structures of the fetal African elephant, Loxodonta africana (Proboscidea, Elephantidae). Acta Zoologica 85, 41-52.
Abstract: The vomeronasal organ (VNO) is a chemosensory structure of the nasal septum found in most tetrapods. Although potential behavioural correlates of VNO function have been shown in two of the three elephant species, its morphology in Loxodonta africana has not been studied. The development of the VNO and its associated structures in the African elephant are described in detail using serially sectioned material from fetal stages. The results show that many components of the VNO complex (e.g. neuroepithelium, receptor-free epithelium, vomeronasal nerve, paravomeronasal ganglia, blood vessels, vomeronasal cartilage) are well developed even in a 154-day-old fetus, in which the VNO opens directly into the oral cavity with only a minute duct present. However, the vomeronasal glands and their ducts associated with the VNO were developed only in the 210-day-old fetus. Notably, in this fetus, the vomeronasal-nasopalatine duct system had acquired a pathway similar to that described in the adult Asian elephant; the VNOs open into the oral cavity via the large palatal parts of the nasopalatine ducts, which are lined by a stratified squamous epithelium. The paired palatal ducts initially coursed anteriorly at an angle of 45degrees from the oral recess and/or the oral cavity mucosa, and merged into the vomeronasal duct. This study confirms the unique characteristics of the elephant VNO, such as its large size, the folded epithelium of the VNO tube, and the dorsomedial position of the neuroepithelium. The palatal position and exclusive communication of the VNO with the oral cavity, as well as the partial reduction of the nasopalatine duct, might be re

Hatfield, J.R., Samuelson, D.A., Lewis, P.A., Chisholm, M., 2004. Structure and presumptive function of the iridocorneal angle of the West Indian manatee (Trichechus manatus), short-finned pilot whale (Globicephala macrorhynchus), hippopotamus (Hippopotamus amphibius), and African elephant (Loxodonta africana). Vet Ophthalmol. 6, 35-43.
Abstract: The iridocorneal angles of prepared eyes from the West Indian manatee, short-finned pilot whale, hippopotamus and African elephant were examined and compared using light microscopy. The manatee and pilot whale demonstrated capacity for a large amount of aqueous outflow, probably as part of a system compensating for lack of ciliary musculature, and possibly also related to environmental changes associated with life at varying depths. The elephant angle displayed many characteristics of large herbivores, but was found to have relatively low capacity for aqueous outflow via both primary and secondary routes. The hippopotamus shared characteristics with both land- and water-dwelling mammals; uveoscleral aqueous outflow may be substantial as in the marine mammals, but the angular aqueous plexus was less extensive and a robust pectinate ligament was present. The angles varied greatly in size and composition among the four species, and most structures were found to be uniquely suited to the habitat of each animal. Department of Small Animal Clinical Sciences, College of Veterinary Medicine, University of Florida, Gainesville, Florida 32610, USA.

Hoffmann, J.N., Montag, A.G., Dominy, N.J., 2004. Meissner corpuscles and somatosensory acuity: the prehensile appendages of primates and elephants
669. Anat. Rec. A Discov. Mol. Cell Evol. Biol. 281, 1138-1147.
Abstract: Meissner corpuscles (MCs) are specialized mechanoreceptors located exclusively in the papillae of glabrous skin. They are confined largely to cutaneous pads of the extremities and respond to transient, phasic, or vibratory stimuli. Though absent in most eutherian taxa, MCs are reported in all primates studied, being most developed in modern humans. The location of MCs between the internal ridges of the epidermis indicates they are well situated to detect friction or deformation at the external surface. Accordingly, MCs are hypothesized to provide primates generally with an enhanced tactile perception. However, the selective pressures favoring greater somatosensory acuity in primates are seldom considered. Interestingly, primate digital dexterity varies greatly. In general, dexterity improves with the extent to which foraging requires food manipulation or textural evaluation. This observation implies that MC density could vary accordingly. Here we report on the density of MCs in five anthropoid taxa selected to represent diverse dietary regimes. Results show that greater MC density correlates with the extent to which primates are frugivorous; however, locomotor and/or phylogenetic effects cannot be discounted

Lazar, J., Rasmussen, L.E., Greenwood, D.R., Bang, I.S., Prestwich, G.D., 2004. Elephant albumin: a multipurpose pheromone shuttle
691. Chem. Biol. 11, 1093-1100.
Abstract: (Z)-7-dodecenyl acetate (Z7-12:Ac) is present in the urine of female Asian elephants (Elephas maximus) approaching ovulation and functions as a female-to-male sex pheromone. Here we show that a significant fraction of the pheromone in the urine is bound to a protein, elephant serum albumin (ESA), and provide evidence for key physiological functions of urinary ESA. Our biochemical and behavioral experiments suggest a three-fold role of ESA in pheromone signaling: (1) transporting Z7-12:Ac from serum into urine; (2) extending the presence of the pheromone in the environment without hampering detection; and (3) targeting pheromone delivery to chemosensory organs through localized release of the ligand induced by a pH change. The exploitation of albumin in pheromone transport clearly distinguishes the elephant from other mammals studied, and complements the uniqueness of elephant anatomy, physiology, and behavior

Leal, W.S., 2004. Pheromone unwrapping by pH flip-flopping
692. Chem. Biol. 11, 1029-1031.
Abstract: The Asian elephant utilizes the same sex pheromone as a number of moth species, (Z)-7-dodecen-1-yl acetate encapsulated in a serum-derived albumin. The chemical signal is emitted in the urine and received in the mucus of the trunk. The unwrapping of the package is pH mediated

Repin, V.E., Taranov, O.S., Ryabchikova, Tikhonov, A.N., Pugachev, V.G., 2004. Sebaceous glands of the woolly mammoth, Mammothus primigenius Blum: histological evidence
651. Dokl. Biol. Sci. 398, 382-384.

Weissengruber, G.E., Forstenpointner, G., 2004. Musculature of the crus and pes of the African elephant (Loxodonta africana): insight into semiplantigrade limb architecture
689. Anat. Embryol. (Berl) 208, 451-461.
Abstract: The limbs of elephants are designed to support the weight of the largest terrestrial animal, and they display unique morphological peculiarities among mammals. In this article we provide a new and detailed anatomical description of the muscles of the lower hindlimb in African elephants (Loxodonta africana), and we place our observations into a comparative anatomical as well as a functional morphological context. At the cranial aspect of the shank (crus) and the foot (pes), the flexors of the tarsal joint and the extensors of the toes form a flat muscular plate covering the skeletal elements. Caudal to the tibia and the fibula the Musculus (M.) soleus is strongly developed, whereas the M. gastrocnemius and the M. flexor digitorum superficialis are thin. Small flexors, adductors, and abductors of the toes are present. The M. tibialis caudalis as well as the Mm. fibularis longus and brevis mainly support the tarsal joint. The design of the muscular structures matches the specific requirements of heavy-weight bearing as well as of proboscidean limb posture and locomotion patterns

2003. Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi.

Allen, W.R., Mathias, S., Wooding, F.B., van Aarde, R.J., 2003. Placentation in the African elephant (Loxodonta africana): II morphological changes in the uterus and placenta throughout gestation. Placenta 24, 598-617.
Abstract: The gross and microscopic development of the zonary endotheliochorial placenta in the African elephant was studied in 22 gravid uteri that ranged in
gestational stage from 0.5 to 20.6 months. The conceptus only ever occupies one horn of the uterus and is associated with 2-5 large corpora lutea that persist in the ipsilateral ovary throughout gestation. Initially, the trophoblast in the equatorial region of the conceptus completely replaces the lumenal epithelium of the endometrium to which it is apposed. Blunt upgrowths of endometrial stroma then develop, each closely invested by trophoblast, and containing the capillaries that will vascularize this maternal component of the resulting placental band. With advancing gestation the lamellate stromal upgrowths increase markedly in length and become much thinner, thereby bringing the trophoblast into intimate contact with the endothelium of the maternal capillaries. They also become folded or pleated to increase the total area of intimate feto-maternal contact. At the lateral edges of the placental band the lamellae bend over towards the endometrium to form a blind cleft. Leakage of blood into this area creates haemophagous zones in which phenotypically specialized trophoblast cells phagocytose the blood components. The presence of large resorbing blood clots and circumferential scars in the uteri of three post parturient animals initiated the hypothesis that, when the standing elephant gives birth at term, the passage of the 120 kg fetus through the vagina may wrench the placenta off the endometrium by severing its very narrow maternal placental hilus. The resulting intrauterine haemorrhage may then play a role in preventing further conception for around 2 years.Department of Clinical Veterinary Medicine Equine Fertility Unit, University of Cambridge, Mertoun Paddocks,Woodditton Road, Suffolk CB8 9BH, Newmarket, UK. vetart@aht.org.uk

Barasa, A., 2003. Morphology and structure of the nictitating membrane cartilage in mammals. Morphologie 87, 5-12.
Abstract: In 30 species of Mammals of varying body size (from rat to elephant), the form, dimensions and structure of the cartilage of the third eyelid were studied. The cartilage is a thin lamina concave in its corneal side, usually elongated in the oro-aboral direction. In the most species studied the outline of the cartilage may be inscribed in a triangle with a oral base, a dorsal margin, a ventral margin and an aboral apex. A study of stained sections revealed, in more than half of species, the presence of elastic fibres in the aboral part of cartilage; these fibres are particularly numerous, but non uniformly distributed, in the Equidae, lion and Suidae. Departement de Morphophysiologie Veterinaire, Rue L. de Vinci 44, 10095 Grugliasco, Italie.

Bhattacharyya, B.K., 2003. Some reproductive traits of elephants. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 37-41.

Clauss, M., Frey, R., Kiefer, B., Lechner-Doll, M., Loehlein, W., Polster, C., Rossner, G.E., Streich, W.J., 2003. The maximum attainable body size of herbivorous mammals: morphophysiological constraints on foregut, and adaptations of hindgut fermenters. Oecologia 136, 14-27.
Abstract: An oft-cited nutritional advantage of large body size is that larger animals have lower relative energy requirements and that, due to their increased gastrointestinal tract (GIT) capacity, they achieve longer ingesta passage rates, which allows them to use forage of lower quality. However, the fermentation of plant material cannot be optimized endlessly; there is a time when plant fibre is totally fermented, and another when energy losses due to methanogenic bacteria become punitive. Therefore, very large herbivores would need to evolve adaptations for a comparative acceleration of ingesta passage. To our knowledge, this phenomenon has not been emphasized in the literature to date. We propose that, among the extant herbivores, elephants, with their comparatively fast passage rate and low digestibility coefficients, are indicators of a trend that allowed even larger hindgut fermenting mammals to exist. The limited existing anatomical data on large hindgut fermenters suggests that both a relative shortening of the GIT, an increase in GIT diameter, and a reduced caecum might contribute to relatively faster ingesta passage; however, more anatomical data is needed to verify these hypotheses. The digestive physiology of large foregut fermenters presents a unique problem: ruminant-and nonruminant-forestomachs were designed to delay ingesta passage, and they limit food intake as a side effect. Therefore, with increasing body size and increasing absolute energy requirements, their relative capacity has to increase in order to compensate for this intake limitation. It seems that the foregut fermenting ungulates did not evolve species in which the intake-limiting effect of the foregut could be reduced, e.g. by special bypass structures, and hence this digestive model imposed an intrinsic body size limit. This limit will be lower the more the natural diet enhances the ingesta retention and hence the intake-limiting effect. Therefore, due to the mechanical characteristics of grass, grazing ruminants cannot become as big as the largest browsing ruminant. Ruminants are not absent from the very large body size classes because their digestive physiology offers no particular advantage, but because their digestive physiology itself intrinsically imposes a body size limit. We suggest that the decreasing ability for colonic water absorption in large grazing ruminants and the largest extant foregut fermenter, the hippopotamus, are an indication of this limit, and are the outcome of the competition of organs for the available space within the abdominal cavity. Our hypotheses are supported by the fossil record on extinct ruminant/tylopod species which did not, with the possible exception of the Sivatheriinae, surpass extant species in maximum body size. In contrast to foregut fermentation, the GIT design of hindgut fermenters allows adaptations for relative passage acceleration, which explains why very large extinct mammalian herbivores are thought to have been hindgut fermenters. Institute of Animal Physiology, Physiological Chemistry and Animal Nutrition, Ludwig-Maximilians-University, Veterinaerstrasse 13, 80539, Munich, Germany. clauss@tiph.vetmed.uni-muenchen.de

Das, D., 2003. Breeding, reproduction and conservation strategies in Asian elephants. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 45-57.

Dutta, D., 2003. Physiology of elephant. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 17-22.

Hutchinson, J.R., Famini, D., Lair, R., Kram, R., 2003. Biomechanics: Are fast-moving elephants really running? Nature 422, 493-494.

Isaza, R., Behnke, B.J., Bailey, J.K., McDonough, P., Gonzalez, N.C., Poole, D.C., 2003. Arterial blood gas control in the upright versus recumbent Asian elephant. Respir Physiolo Neurobiol 134, 169-176.
Abstract: In the elephant, there is concern that lateral recumbency (LR) impairs respiratory muscle and lung function resulting in clinically significant arterial hypoxemia. Using healthy adult female Asian elephants (Elephas maximus, n=6), the hypothesis was tested that, given the O₂ binding characteristics of elephant blood, substantial reductions in arterial O₂ pressure PaO₂ in LR could be tolerated without lowering arterial O₂ content appreciably. Fifteen minutes of LR decreased PaO₂ from 103+/-2 (upright, U) to 77+/-4 mmHg (P<0.05) and hemoglobin O₂ saturation (U, 97.8+/-0.1, LR, 95.3+/-0.5%, P<0.05). However, due to a recumbency-induced hemoconcentration, arterial O₂ content was unchanged (U, 18.2+/-2.4, LR, 18.3+/-2.1 ml O₂ per 100 ml). In addition, there was a mild hyperventilation in LR that reduced arterial CO₂ pressure (PCO₂₎ from 39.4+/-0.3 to 37.1+/-1.0 mmHg (P<0.05). These data indicate that the Asian elephant can endure at least short periods of LR without lowering arterial O₂ content.

Kalita, S.N., Sarma, M., 2003. Anatomy of elephant: some important features. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 10-16.

Nalla, R.K., Kinney, J.H., Ritchie, R.O., 2003. Effect of orientation on the in vitro fracture toughness of dentin: the role of toughening mechanisms. Biomaterials 24, 3955-3968.
Abstract: Toughening mechanisms based on the presence of collagen fibrils have long been proposed for mineralized biological tissues like bone and dentin; however, no direct evidence for their precise role has ever been provided. Furthermore, although the anisotropy of mechanical properties of dentin with respect to orientation has been suggested in the literature, accurate measurements to support the effect of orientation on the fracture toughness of dentin are not available. To address these issues, the in vitro fracture toughness of dentin, extracted from elephant tusk, has been characterized using fatigue-precracked compact-tension specimens tested in Hank's balanced salt solution at ambient temperature, with fracture paths perpendicular and parallel to the tubule orientations (and orientations in between) specifically being evaluated. It was found that the fracture toughness was lower where cracking occurred in the plane of the collagen fibers, as compared to crack paths perpendicular to the fibers. The origins of this effect on the toughness of dentin are discussed primarily in terms of the salient toughening mechanisms active in this material; specifically, the role of crack bridging, both from uncracked ligaments and by individual collagen fibrils, is considered. Estimates for the contributions from each of these mechanisms are provided from theoretical models available in the literature. Materials Sciences Division, Lawrence Berkeley National Laboratory, Department of Materials Science and Engineering, University of California, Berkeley, CA 94720, USA.

Pothiwong, W., Kamonrat, P., Uthaichotiwan, P., 2003. A morphological study and diagnotic ultrasonography of Asian elephant kidney. Thai Journal of Veterinary Medicine 33, 79-88.

Rajaram, A., Krishnamurthy, V., 2003. Elephant temporal gland ultrastructure and androgen secretion during musth. Current Science 85, 1467-1471.
Abstract: We have investigated the ultrastructure of the temporal gland of the Asian elephant (Elephas maximus) in the musth condition. We find that the organelles are highly evolved for the production of the androgen, testosterone which is reported to be very high in the Asian male elephant in full musth. The mitochondria bear cristae which are profuse and tubular, and occur along with many Golgi bodies. There is hypertrophy of smooth endoplasmic reticulum. All the structures involved in the production of androgen, as in the Leydig cell or the cells of the adrenal cortex, are thus found in abundance. Cellular structures also seem singularly evolved for the secretion of androgen and its degradation products.

Steenkamp, G., 2003. Oral biology and disorders of tusked mammals. Veterinary Clin North Am Exot Anim Pract. 6, 689-725.
Abstract: Tusked mammals can be terrestrial or aquatic. Many of these magnificent animals are kept in captivity all over the world. Functions of tusks vary as much as the species in which they occur. Dental anomalies and disorders of tusks and the rest of the dentition in these mammals were discussed, with an emphasis on the elephant. The tusk anatomy, with its large, conically-shaped pulp, makes it an ideal tooth for partial pulpectomy treatment in trauma cases where the pulp is exposed. Surgical techniques for tusks have been developed and were discussed. Oral tumors occur, but are rare.Department of Companion Animal Clinical Studies, Faculty of Veterinary Science, University of Pretoria, Private Bag X04, Onderstepoort 0010, South Africa. steenkamp@op.up.ac.za

West, J.B., Fu, Z., Gaeth, A.P., Short, R.V., 2003. Fetal lung development in the elephant reflects the adaptations required for snorkeling in adult life. Respir Physiol Neurobiol 138, 325-333.
Abstract: The adult elephant is unique among mammals in that the pleural membranes are thickened and the pleural cavity is obliterated by connective tissue. It has been suggested that this peculiar anatomy developed because the animal can snorkel at depth, and this behavior subjects the microvessels in the parietal pleura to a very large transmural pressure. To investigate the development of the parietal pleura, the thickness of the endothoracic fascia (ET) was measured in four fetal African elephants of approximate gestational age 111-130 days, and the appearances were compared with those in human, rabbit, rat and mouse fetuses of approximately the same stage of lung organogenesis. The mean thicknesses of ET in the elephant, human, rabbit, rat and mouse were 403, 53, 29, 27 and 37 microm, respectively. This very early development of a thick parietal pleura in the elephant fetus is consistent with the hypothesis of a long history of snorkeling in the elephant's putative aquatic ancestors. Department of Medicine, University of California San Diego, 9500 Gilman Drive, La Jolla, CA 92093-0623, USA. jwest@ucsd.edu

Yappert, M.C., Rujoi, M., Borchman, D., Vorobyov, I., Estrada, R., 2003. Glycero- versus sphingo-phospholipids: correlations with human and non-human mammalian lens growth. Exp Eye Res 76, 725-734.
Abstract: The human lens differs from other mammalian lenses in its very slow growth and unusual phospholipid composition of its cell membranes. Dihydrosphingomyelins (DHSMs) make up about half of all phospholipids in adult human fiber membranes. In all other membranes, sphingomyelins(SMs) with a trans double bond in their backbone, are prevalent. In our quest to understand the biological implications of such elevated DHSM levels, we analyzed membranes from various regions of human, elephant, giraffe, polar bear, pig and cow lenses. The levels of DHSMs were minor in non-human lens membranes. A strong correlation was observed between growth rate and relative contents of phosphatidylcholines(PCs) in epithelia and outer cortical fibers. Sphingomyelins became increasingly predominant in differentiated fibers and this increase was age dependent. Indeed, nuclear fiber membranes of aged non-human mammals were composed, almost exclusively, of (SMs). Although human lens membranes followed comparable compositional trends, the magnitude of the changes was much smaller. We postulate that the high relative contents of DHSMs provide a biochemically inert matrix in which only small amounts of PCs and SMs and their metabolites, known to promote and arrest growth, respectively, are present. This compositional difference is proposed to contribute to the slow multiplication and elongation of human lens cells. Department of Chemistry, College of Arts and Sciences, University of Louisville, Louisville, KY 40292, USA. mcyapper@louisville.edu

Allen, W.R., Mathias, S.S., Wooding, F.B., Skidmore, J.A., van Aarde, R.J., 2002. Placentation in the African elephant, Loxodonta africana. I. Endocrinological aspects. Reprod Suppl 60, 105-116.
Abstract: Placental and fetal tissues were recovered from the uteri of 59 pregnant elephant that ranged in estimated age from day 18 to month 21 of gestation. Incubation of placenta and fetal gonad, alone or in combination, with tritium-labelled cholesterol, pregnenolone and androstenedione failed to yield any labelled progestagens or oestrogens from placenta, but did produce small amounts of labelled progesterone and 5alpha-dihydroprogesterone from fetal gonad. Immunochemical staining of tissues with four antisera specific for enzymes involved in the steroidogenic pathway revealed no staining in sections of placenta but positive labelling for P450 side chain cleavage enzyme (SCC450) and 3beta-hydroxysteroid dehydrogenase (3beta-HSD) of the interstitial cells that comprise the bulk of the enlarged fetal gonads during the second half of gestation. Saline extracts of placental tissue showed no activity in three different gonadotrophin assays. In view of this endocrinological inactivity in the zonary elephant placenta and the probable reliance on maternal luteal sources of progestagens for maintenance of the pregnant state, the argument is advanced that uncomplicated abortion would probably follow a single administration of a PGF analogue given at any stage of pregnancy. If so, the treatment might constitute an efficacious method for controlling population increases in elephants maintained in enclosed game parks in Africa.

Chandrasekharan, K., 2002. Elephant - an overview. Journal of Indian Veterinary Association Kerala 7, 8-11.

Cheeran, J.V., 2002. Elephant facts. Journal of Indian Veterinary Association Kerala 7, 12-14.

Endo, H., Akihisa, N., Sasaki, M., Yamamoto, M., Arishima, K., 2002. The renal structure in an Asian elephant (Elephas maximus). Anat Histol Embryol 31, 269-272.
Abstract: The renal structure of a female Asian elephant (Elephas maximus) was observed in both macroscopic and light microscopic levels. The left kidney was elongated-ellipse in shape, whereas the right appeared round. The left kidney was 31 cm in cranio-caudal length, 21 cm in medio-lateral length, and 2950 g in weight. The right kidney was 34 cm in cranio-caudal length, 22 cm in medio-lateral length, and 3250 g in weight. The external appearance showed the six separated renal lobes in both sides of the kidney. The four pairs of the lobes were fused in the deepest region in both sides of kidney, so we considered it as an incompletely lobated kidney in this species. We observed the proximal and distal urinary tubules in histological sections. Many renal corpuscles consisted of the glomerulus and Bowman's capsule. Many mesangial cells and some podocytes were confirmed in each glomerulus; however, Bowman's capsules were larger than those in other mammalian species.

Johnson, E.W., Rasmussen, L., 2002. Morphological characteristics of the vomeronasal organ of the newborn Asian elephant (Elephas maximus). Anatomical Record 267, 252-259.
Abstract: The 6-week-old Asian elephant (Elephas maximus) has a well-documented precocious flehmen response to pheromones, suggesting that the pheromone-detecting vomeronasal organ (VNO) is functional very early in the life of this species. To further document this, the VNOs of two newborn elephants were examined in situ and analyzed by light microscopy (LM) to ascertain their structural maturity at birth. A tubular, cartilage-encased VNO was located along the anterior base of each side of the nasal septum. Its rostral end was connected to a duct to the roof of the mouth; the caudal end was attached to a well-defined vomeronasal nerve projecting toward the brain. LM revealed distinctive differences in the mucosae bordering the horseshoe-shaped lumen: a concave, sensory mucosa, and a convex, nonsensory mucosa. Small groups of receptor neurons were observed among ciliated columnar cells in the sensory epithelium. Numerous unmyelinated nerve bundles and blood vessels filled the underlying lamina propria (LP) and a small section of the vomeronasal nerve was conspicuous at one edge. The nonsensory mucosa manifested a thinner epithelium that principally consisted of ciliated columnar cells, some of which showed a granular cytoplasm, and a conspicuous row of basal cells. The LP was replete with acinar glands and ducts that opened into the lumen. This study shows that the VNO of the newborn elephant has reached an advanced stage of structural maturity, closely resembling that of the adult. Its composition supports the view that flehmen at 6 weeks delivers pheromones to a functional VNO.

Kim, C.S., Won, C.K., Cho, G.H., Cho, K.W., Park, J.S., Rho, G.J., 2002. A case of fused thoracic vertebrae, and lumbar vertebrae, sacrum and ilium of African elephant (Loxodonta africana). Korean Journal of Veterinary Research 42, 131-136.

Kumar, G.A., Ghosh, K.N.A., Sreekumaran, T., Chandrasekharan, K., 2002. Reproduction in elephants. Journal of Indian Veterinary Association Kerala 7, 38-40, 48-48.

Langman, V.A., Rowe, M., Maloney, M., McQuire, R., Carrington, R. Obligatory heterothermia a story of elephant radiators. Proceedings of the Elephant Managers Association Conference. Journal of Experimental Biology, Lond. 22, 136. 2002.
Ref Type: Conference Proceeding

Rees, P.A., 2002. Asian elephants (Elephas maximus) dust bathe in response to an increase in environmental temperature. Journal of Thermal Biology 27, 353-358.
Abstract: (1) A captive herd of Asian elephants (Elephas maximus) exhibited dusting behaviour when the maximum daily temperature exceeded approximately 13°C, and dusting frequency increased directly with the environmental temperature. (2) Individual animals showed variation in dusting frequency but this was not related to body mass, suggesting that the function of dusting is not primarily thermoregulatory. (3) Synchronisation in the timing of dusting behaviour within the herd suggests that it may have a function in the maintenance of social cohesion. (4) The function of dusting behaviour could not be determined from the data presented, but it may be involved in skin care, protection from insects or other parasites, temperature control, protection from radiation or some combination of these.

Saseendran, P.C., Anil, K.S., Nair, A., Radhakrishnan, K., Prasad, A., 2002. Elephants and work. Journal of Indian Veterinary Association Kerala 7, 41, 48-48.

Soltysiak, Z., Barcikowska, M., Nieman, S., 2002. Specification patterns of amyloid-beta deposits in old fish, reptile, bird and several old mammal brains. Medycyna \Weterynaryjna 58, 74-76.
Abstract: The study was conducted to demonstrate whether there are amyloid-beta deposits in old fish, reptiles, birds and some mammals. The thick brain sections were stained with cresyl violet, Congo red, tioflavin S and by immunocytochemistry using mAb 4G8. Amyloid-beta deposits were not found in old fish, reptile and bird brains. Amyloid-beta was found in old mammal brains in three forms: parenchymal amyloidosis, congophilic angiopathy and diffuse amyloid deposits. Three types of plaque were found in old mammal brain cortexes: diffuse, primitive, and classic (neuritic). The neuritic plaque alveas consisted of three components: degenerating neurites, amyloid, and reactive cells. Parenchymal amyloidosis and congophilic angiopathy were found only in five mammal brains, two from wolf, two from fox and one from elephant brain. Congophilic angiopathy was discovered in all investigated mammals.

Sreekumar, K.P., 2002. Physiological features of Indian elephant. Journal of Indian Veterinary Association Kerala 7, 35-36.

West, J.B., 2002. Why Doesn't the Elephant Have a Pleural Space? News Physiol Sci 17, 47-50.
Abstract: The elephant is the only mammal whose pleural space is obliterated by connective tissue. This has been known for 300 years but never explained. The elephant is also the only animal that can snorkel at depth. The resulting pressure differences require changes in the pleural membranes and pleural space.

Witter, K., Matulova, P., Misek, I., 2002. The lateral enamel lamina--component of tooth primordia in selected mammalian species. Connect Tissue Res 42, 134-137.
Abstract: The lateral enamel lamina (LEL) is a part of the enamel organ, which is probably not involved in tooth formation. It represents, besides the "stalk" of the tooth primordium, a second interconnection between enamel organ and oral epithelium or vestibular lamina. We detected the LEL in the sheep (Ovis aries), the dolphin (Stenella attenuata), and the vole (Microtus agrestis) by light microscopy and computer-aided three-dimensional reconstruction. The LEL could be found in cap to bell stage tooth primordia, most clearly in slowly developing tooth germs. LEL-like structures have been furthermore described or depicted in tooth germs of the mouse, the elk (Alces alces), the dugong (Dugong dugong), the elephant (Loxodonta africana), and the human. Probably it is a part of all mammalian tooth primordia that undergoes regression during morphogenesis of the enamel organ. As a reducing structure, it should be considered in studies of tooth development.

Ball, R.L. Ultrasound Evaluation of the Pleura Space and Associated Connective Tissue in the Asian Elephant (Elephas maximus). A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001. 245. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Cozzi, B., Spagnoli, S., Bruno, L., 2001. An overview of the central nervous system of the elephant through a critical appraisal of the literature published in the XIX and XX centuries. Brain Res Bull 54, 219-227.
Abstract: The two species of elephants (Indian: Elephas maximus and African: Loxodonta africana) possess the largest brain among land mammals. Due to its size, the elephant brain is discussed in virtually every paper dealing with the evolution of the central nervous system of mammals and comparative brain size. Studies on the social habits of elephants also deal with the skills and the "intelligence" and brain size of these species. Yet most of the descriptions and conclusions reported in comparative studies rely on second-hand data derived from investigations performed several decades before, often dating as far back as the XIX century. Furthermore, many of the original papers actually describing gross and detailed features of the brain of elephants are either no longer available, are written in languages other than English, or are difficult to trace. The present study gives a short description of the anatomy of the central nervous system of elephants, with special attention to its distinctive features, reports all available literature on the subject, and briefly discusses its origins and rationale.

Dehnhard, M., Hildebrand, T., Rohleder, M., Strauss, G., Meyer, H.H.D., Goritz, F., 2001. Application of an enzyme-immunoassay (EIA) for rapid screening of 5alpha-pregnane-3,20-dione (DHP) in blood plasma of the Asian elephant, Elephas maximus. Berliner und Munchener Tierarztliche Wochenschrift. 114, 161-165.
Abstract: Populations of African (Loxodonta africana) and Asian elephants (Elephas maximus) in zoos and safari parks are at risk due to their low reproductive success. To extend the limited knowledge of their reproductive physiology, easy and practical methods for the analysis of relevant reproductive hormones must be developed to support assisted reproduction, for instance. Blood samples from 2 nonpregnant and 2 pregnant Asian elephants were used in the following study. For the measurement of 5alpha-pregnane-3,20-dione (DHP), the predominant ovarian gestagen in both species, an enzyme-immunoassay (EIA) based on commercial reagents was applied. Advantages of this EIA are the small volume of plasma needed for evaluation (5 µl) and the possibility of direct processing without an extraction stage. The lower limit of detection was 0.16 ng/ml, mean recovery was 101% and the mean coefficients of variation were 7.3 (intra-assay) and 9.9% (inter-assay). In the Asian elephants, DHP levels reached 15 ng/ml during the luteal phase and 21 ng/ml during pregnancy. Oestrous cycle lengths based on the lowest DHP concentrations varied from 12 to 20 weeks (mean of 15.4±2.3). In two Asian elephant cows, a calf was stillborn. Thereafter, ovarian activity in the animals resumed after approximately 8 and 13 weeks, respectively. In one animal, estradiol implants for hormonal contraception caused a down regulation of ovarian function as demonstrated by an irregular pattern of DHP secretion over a period of 48 weeks. It is proposed that the direct DHP-EIA is a suitable method for reproductive monitoring in elephants, as it can be easily established in laboratories.

Endo, H., Hayashi, Y., Komiya, T., Narushima, E., Sasaki, M., 2001. Muscle architecture of the elongated nose in the Asian elephant (Elephas maximus). Journal of Veterinary Medical Science 63, 533-537.
Abstract: The architecture of the M. caninus in the elongated nose was examined in the Asian elephant (Elephas maximus). The following complicated musculature of the M. caninus was observed in the proximal and distal regions of the nose: (1) Proximal region: In the superficial layer, the longitudinal bundles are confirmed in the dorsal part, and the obliquely-oriented ones in the ventral part. In the middle layer, some bundles run ventro-distally, while other ones represent longitudinally-oriented running. The deep layer consists of complicated architecture of many bundles. Some muscle bundles run medio-laterally, while the others extend proximo-distally in this space. (2) Distal region: In the dorsal part of the M. caninus, the bundles run at deep-superficial direction, while in the ventral part the bundles are longitudinally arranged. The bundles run at lateral direction near the septum of the nasal conduits. The N. facialis and N. infraorbitalis send many branches in the lateral area of the M. caninus in the trunk. This muscle architecture of multi-oriented bundles and well-developed innervation to them suggest that they enable the elongated nose to act as a refined manipulator in the Asian elephant.

Ganslober, U. Behavioural Ecology, Social Relationships, Life History and Evolutionary Constraints in Megaherbivores. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001. 26-31. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Hinke, A., Wipplinger, J. A Case of Molar Anomalie in an Asian Elephant (Elephas maximus). A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001. 264. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Lamps, L.W., Smoller, B.R., Rasmussen, L.E., Slade, B.E., Fritsch, G., Goodwin, T.E., 2001. Characterization of interdigital glands in the Asian elephant (Elephas maximus). Research in Veterinary Science 71, 197-200.
Abstract: In the Asian elephant, wetness akin to perspiration is commonly observed on the cuticles and interdigital areas of the feet; this observation has lead to speculation regarding the existence of an interdigital gland. Our goal was to search for interdigital glands and characterize them morphologically, histochemically, and immunohistochemically. Necropsy samples of interdigital areas from two Asian elephants were obtained. Multiple sections were fixed and processed routinely, then stained with hematoxylin/eosin and differential mucin stains. Immunohistochemistry was also performed for cytokeratins 8 and 10. Interdigital glands resembling human eccrine glands were detected deep within the reticular dermis. Histochemical staining indicated neutral mucopolysaccharides and nonsulphated acid mucopolysaccharides in glandular secretions, and the glandular epithelium also showed immunoreactivity to cytokeratins 8 and 10. Both the histochemical and immunohistochemical staining patterns are analogous to human eccrine structures. This study shows with certainty that Asian elephants possess sweat glands as they are defined histologically.

Phillips, P.K., Heath, J.E., 2001. Heat loss in Dumbo: a theoretical approach. Heat loss in Dumbo: a theoretical approach 26, 117-120.
Abstract: A flat plate model was used to calculate heat loss from the pinnae of the animated elephant Dumbo. In conditions of high wind velocity and large
gradients, Dumbo could potentially dissipate more heat than he produces. This suggests that he may need the large ears to help lose the excess heat produced while flying.

Ramsay, E., Henry, R., 2001. Anatomy of the Elephant Foot. In: Csuti, B., Sargent, E.L., Bechert, U.S. (Eds.), The Elephant's Foot. Iowa State University Press, Ames IA, USA, pp. 9-12.
Abstract: While elephants have played an important role in human ecology throughout recorded history, their anatomy has been the subject of relatively few studies, and these studies considered only a limited number of specimens. This is especially true for the elephants' feet, despite the common occurrence of foot disease in elephants (Evans 1961, Mikota et al. 1994). One study that surveyed North American captive elephants found that 50 percent had experience foot problems (Mikota et al. 1994). Despite their phylogenetic differences, the components of Asian (Elephas maximus) and African (Loxodonta africana) elephants feet are remarkably similar. The major differences are the shape of the rear foot and the number of phalanges and toenails. This is not to say that the feet of the two species are the same. The differences in their wild habitats and the more common occurrence of foot lesions in captive Asian elephants suggest that the biology of the two species' feet are quite different. The following describes anatomical characteristics common to both species unless otherwise identified.

Singer, M.A., 2001. Of mice and men and elephants: metabolic rate sets glomerular filtration rate. Am J Kidney Dis 37, 164-178.
Abstract: Allometric scaling deals with the functional consequences of changes in size or scale among geometrically dissimilar animals (ie, animals differing in proportions). For adult mammals ranging in size and proportion from mouse to elephant, the data describe an interdependent set of functions consisting of metabolism (measured as metabolic rate), glomerular filtration rate (GFR), effective renal plasma flow, excretion of nitrogenous waste products, cardiac output, and pulmonary function-related variables. Within this set of functions, evidence indicates that metabolic rate is the primary process. One important design feature is given by the ratio of GFR to metabolic rate. Because this ratio is independent of size, it can be generalized to all mammals in this series. The numeric value of this ratio gives the optimal GFR for each unit of metabolic rate. A simple hypothesis is proposed: metabolic rate, the primary process, sets GFR. This relationship is unidirectional. A decrease in GFR, for example, caused by nephron loss, should not lead to a change in metabolic rate. This hypothesis was tested in four natural experiments: human growth and development, thyroid dysfunction, chronic renal failure, and hibernation. The results are consistent with this hypothesis.

West, J.B., 2001. Snorkel breathing in the elephant explains the unique anatomy of its pleura. Respiratory Physiology 126, 1-8.
Abstract: It has been known for over 300 years that the anatomy of the elephant lung is unique among mammals in that the pleural cavity is obliterated by connective tissue. However no satisfactory explanation has been advanced. Recent studies suggest that the elephant has an aquatic ancestry and the trunk may have developed for snorkeling. In addition, the modern day elephant is the only mammal that can remain submerged far below the surface of the water while snorkeling. The resulting differences of pressures within the thorax mean that the small blood vessels of the pleura are in great danger of rupturing or causing severe edema. The same distribution of pressures occurs when the animal raises water inside its trunk prior to drinking although in this case the pressure differences are relatively short-lived. Evolution has provided a remarkable solution to this problem by replacing the normally delicate parietal and visceral pleurae by dense connective tissue, and separating the two pleurae by loose connective tissue to allow some sliding movement.

Crossley, D.A., 2000. Elephant tusks: where are the nerves? J. Vet. Dent. 17, 37.

Fritsch, G., Göritz, F., Hermes, R., Jewgenow, K., Maltzan, J., Hildebrandt, T.B., 2000. Physiology of sexual maturity in male elephants. Reprod Dom Anim 35, 26.

Kitiyanant, Y., Schmidt, M.J., Pavasuthipaisit, K., 2000. Evaluation of sperm acrosome reaction in the Asiatic elephant. Theriogenology 53, 887-896.
Abstract: This study focuses on the effect of chemicals on acrosome reaction in elephant spermatozoa. Semen was collected at the Washington Park Zoo in Portland, Oregon, from an 11-yr-old Asian elephant by artificial vagina (7 ejaculates) and transported to Mahidol University in Bangkok in extender at 4 to 5 degrees C within 24 to 28 h. A total of 500 x 10(6) sperm/mL was used for the control and for each of the 4 treatment groups: 1) cAMP (0.1 mM); 2) caffeine (0.1 mM); 3) Penicillamine hypotaurine and epinephrine, PHE (penicillamine 2 mM, hypotaurine 1 mM, epinephrine 1 mM); and 4) heparin (10 microg/mL) at 39 degrees C for 2 h. Aliquots were removed and the sperm viability, abnormal morphology, and acrosome status were evaluated by triple stain technique. Transmission electron microscopy (TEM) was used to observe changes of the sperm head membrane in all treatment groups. Trypan blue reliably stained dead spermatozoa, while rose Bengal stained only the spermatozoa with intact acrosomes. The concentration of dead sperm cells was similar in the 4 groups. The percentages of live acrosome-reacted spermatozoa in the control and in groups treated with caffeine, PHE, cAMP and heparin were 19.5 +/- 4.3, 38.1 +/- 4.0, 34.8 +/- 3.7, 29.8 +/- 0.8 and 28.0 +/- 4.2, respectively. The acrosome reaction rate was higher in the treatment groups than in the control (P<0.05). Caffeine and PHE caused significantly higher acrosome reaction of the sperm head than cAMP or heparin (P<0.05). The electron micrographs showed that the acrosome reaction occurred by the presence of apical vesiculation. The results indicated that 1) the triple stain technique allowed for evaluation of both viability and acrosome reaction simultaneously in elephant spermatozoa,2) acrosome reaction occurred at a high rate in all 3 treatment groups. 3) the effects of caffeine and PHE were significantly higher (P<0.05) than of cAMP and heparin, and 4) the data obtained from the triple stain technique corresponded to those from TEM.

Luikart, K. Anatomy of the Elephant Forefoot. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop. 14-15. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Malik, M.R., Shrivastav, A.B., Jain, N.K., Vaish, R., 2000. Morphometry of kidney of elephant. Indian Journal of Veterinary Anatomy 12 , 101-102.

Malik, M.R., Shrivastava, A.B., Jain, V.K., Rakhi, V., 2000. Lobar pattern of kidneys of elephant. Indian Journal of Veterinary Anatomy 12, 18-22.

Malik, M.R., Shrivastava, A.B., Jain, V.K., Rakhi, V., 2000. Biometry of the cerebral ventricles of elephant. Indian Journal of Veterinary Anatomy 12, 110-112.

Malik, M.R., Shrivastav, A.B., Jain, V.K., Rakhi, V., 2000. A note on encephalometry of Asian elephant. Indian Journal of Veterinary Anatomy 12, 103-104.

Raubenheimer, E.J., 2000. Early development of the tush and the tusk of the African elephant (Loxodonta africana). Arch Oral Biol 45, 983-986.

Schmitt, D.L., Pace, L.W. Multiple Congenital Cardiac Anomalies in a Newborn Asian Elephant (Elephas maximus). Proceedings of the Elephant Managers Association Conference, Oct 6-9,2000 Syracuse, NY. 13-14. 2000. 2000.
Ref Type: Conference Proceeding
Abstract: Cardiac anomalies in humans occur in about 1% of human births. Most are a developmental disorder of the vascular trunk and septum of the heart, which result in reduced blood circulation to periphery. This report of a cardiac anomaly in a neonatal elephant is first to the author's knowledge. A congenital defect known as tetrology of Fallot is described in a male Asian elephant who lived for 9 hours following birth.

Singer, M.A., Morton, A.R., 2000. Mouse to elephant: biological scaling and Kt/V. Am J Kidney Dis 36, 306-309.
Abstract: The construct Kt/V is used by the nephrology community in prescribing dialysis dose. The concerns that have been raised as to what value of V to use in the calculation of Kt/V touch on the more central question of whether filtration rate should be normalized by a parameter other than V. Within the animal kingdom, a number of physiological variables scale to body size according to an equation of the form Y = YoMb, where Yo is a constant, M is body mass, and b is a scaling exponent. Glomerular filtration rate (GFR) in mammals weighing from 30 g to 503 kg scales to body weight with an exponent of 0.77. Hence, GFR per unit body weight (or Kt/V) decreases significantly with increasing body size. Metabolic rate also scales to body size in a wide range of mammals according to the same general equation and with a scaling exponent of 0.75. Because GFR and metabolic rate scale to body mass with virtually the same exponent, a ratio of the two yields a constant independent of body size. We propose that the ratio (filtration rate/metabolic rate) replace Kt/V. Such a ratio would underscore the linkage between filtration rate (and dialysis therapy) and the metabolic demands of the body.

Valli, V.E., Jacobs, R.M., 2000. Structure and function of the hemopoietic system. In: Feldman, B., Zinkl, J.G., Jain, N.C. (Eds.), Schalm's Veterinary Hematology. Lippinicott, Williams & Wilkins, Baltimore, Maryland, USA, pp. 225-239.

Victor, S., Nayak, V.M., 2000. Evolutionary anticipation of the human heart. Ann R Coll Surg Engl 82, 297-302.
Abstract: We have studied the comparative anatomy of hearts from fish, frog, turtle, snake, crocodile, birds (duck, chicken, quail), mammals (elephant, dolphin, sheep, goat, ox, baboon, wallaby, mouse, rabbit, possum, echidna) and man. The findings were analysed with respect to the mechanism of evolution of the heart.

Weissengruber, G.E., Kubber-Heiss, A., Forstenpointner, G., Riccaboni, P., 2000. On the morphology of the temporal gland (Glandula temporalis) in the African elephant (Loxodonta africana). Wiener Tierarztliche Monatsschrift 87, 303-308.

1999. Equine Medicine and Surgery. Mosby, St. Louis MO USA.

Fagan, D.A., Benirschke, K., Simon, J.H., Roocroft, A., 1999. Elephant dental pulp tissue: where are the nerves? J Vet Dent 16, 169-172.
Abstract: Dental pulp tissue from three elephants was examined histologically with hematoxylin and eosin and s-100 protein stains. In all specimens, normal pulp was found with the exception that no nerve fibers (myelinated or non-myelinated) were demonstrable in any of the numerous sections prepared.

Gaeth, A.P., Short, R.V., Renfree, M.B., 1999. The developing renal, reproductive, and respiratory systems of the African elephant suggest an aquatic ancestry. Proc Natl Acad Sci U S A 96, 5555-5558.
Abstract: The early embryology of the elephant has never been studied before. We have obtained a rare series of African elephant (Loxodonta africana) embryos and fetuses ranging in weight from 0.04 to 18.5 g, estimated gestational ages 58-166 days (duration of gestation is approximately 660 days). Nephrostomes, a feature of aquatic vertebrates, were found in the mesonephric kidneys at all stages of development whereas they have never been recorded in the mesonephric kidneys of other viviparous mammals. The trunk was well developed even in the earliest fetus. The testes were intra-abdominal, and there was no evidence of a gubernaculum, pampiniform plexus, processus vaginalis, or a scrotum, confirming that the elephant, like the dugong, is one of the few primary testicond mammals. The paleontological evidence suggests that the elephant's ancestors were aquatic, and recent immunological and molecular evidence shows an extremely close affinity between present-day elephants and the aquatic Sirenia (dugong and manatees). The evidence from our embryological study of the elephant also suggests that it evolved from an aquatic mammal.

Rasmussen, L.E.L., Perrin, T.E., 1999. Physiological correlates of musth: lipid metabolites and chemical composition of exudates. Physiology and Behavior 67, 539-549.
Abstract: Physiological changes related to lipid metabolism, behaviour and chemicals released in body exudates were studied during musth in the Asian elephant (Elephas maximus) as a case study. During musth, changes in serum testosterone and triglyceride concentrations followed similar patterns, with the former increasing sooner than the latter. Deviant behaviour increased during changing androgen levels. The observed high concentrations of testosterone were positively and significantly correlated with increased triglycerides. Lipase activity elevated significantly immediately before and after musth. Blood pH increased significantly in alkalinity. Urine and temporal gland secretions released variable amounts of compounds, some of which may be chemical signals. During musth, temporal gland and urinary exudates demonstrated increased acetone and other ketones indicative of lipid metabolic alterations. Large quantities of nonmethane hydrocarbons, especially 2-butanone, were released from the seemingly dry orifice of the temporal gland before the start of over musth and before maximum blood elevations were observed; isoprene release was similar. However, maximal acetone levels occurred simultaneously in blood, temporal gland secretions, and urine. Metabolically, musth is a series of interwoven, changing stages of increasing and decreasing hormones and lipid-related constituents. Released chemicals can be quantitatively related to these internal physiological events; some observed behaviours appear to result from altered chemical signals.

Silver, W., 1999. Chemesthesis: The burning questions. Chemosense 2.

Abou-Madi, N., Kollias, G.V., Sturmer, A.T., Hackett, R.P. Umbilical herniorrhaphy in a juvenile Asian elephant (Elephas maximus). Proceedings AAZV and AAWV Joint Conference. 212-216. 1998.
Ref Type: Conference Proceeding

Godagama, W.K., Wemmer, C., Ratnasooriya, W.D., 1998. The body condition of Sri Lankan domesticated elephants (Elephas maximus maximus). Ceylon Journal of Science, Biological Sciences 26, 1-5.
Abstract: The objective of this study was to evaluate the body condition of domesticated elephants in Sri Lanka using an index based upon visual assessment and numerical scoring of 6 criteria (temporal depression, scapula, thoracic region, flank area, lumbar vertebrae and pelvic bone) resulting in a scale of 0-11. The study was conducted between 1 April 1993 and 1 April 1994 in 13 administrative districts using 140 domesticated elephants. The mean body condition index of the elephants was 6.95±0.26 points. Males had significantly lower body condition index (6.63±0.22 points) than females (7.3±0.21 points) and there was no significant correlation between age and body condition index. The index was not significantly different between elephants which were owned by private individuals or temples and dewales and maintained by mahouts or their owners.

Godagama, W.K., Wemmer, C., Ratnasooriya, W.D., 1998. Spinal conformation of domesticated Sri Lankan elephants (Elephas maximus maximus). Ceylon Journal of Science, Biological Sciences 26, 7-11.
Abstract: The study investigated whether the 5 spinal conformations previously described for the Burmese elephant (Elephas maximus birmanicus) are also present in the Sri Lankan elephant. 140 domesticated elephants were examined according to Gale's five-description system. The 5 spinal conformation types described for the Burmese elephant are also present in the Sri Lankan elephant. Out of the 140 elephants, 23 (16%) had Type 1, 48 (34%) had Type 2, 5 (4%) had Type 3, 50 (36%) had Type 4 and 14 (10%) had Type 5 spinal conformation. There was a significant variation in the spinal conformation of male and female elephants.

Patil, V.A., Bhamburkar, V.R., Dalvi, R.S., Banubakode, S.B., Kale, M.A., 1998. Morphometrical study of pelvis in some animals. Journal of Bombay Veterinary College 6, 45-46.
Abstract: The morphometric study of the pelvis in buffaloes, cattle, goats, horses, pigs, dogs, panthers, sambar and elephants was carried out. The pelvic index, obturator foramen index, greater sciatic notch index and lesser sciatic notch index were calculated. It is concluded that these indices are useful in species differentiation.

Rasmussen, L.E.L., 1998. Chemical communication: An integral part of functional Asian elephant (Elephas maximus) society. Ecoscience 5, 410-426.
Abstract: The matriarchally organized Asian elephant society is characterized by long-term stability and continuity. Flux within this society results from changing ecological conditions and the dynamics of its population. Its structure is influenced by age composition and physiological states within the female herd and by impinging influences of the peripheral males, especially during reproductive times. Recent behavioral studies of captive populations have substantiated older field studies and have demonstrated that chemical signals play a significant role in elephant society. Chemical investigations, based on previously substantiated behavioral interactions, have identified specific compounds or combinations of compounds in elephant emissions (especially urine, temporal gland secretions and breath) that retain bioactivity throughout chemical extractions and playback experiments, based on behavioral and/or chemosensory responses. Chemosensory neuroreceptive systems in Asian elephants are reviewed, as well as behavioral and chemosensory effects of whole exudate chemical signals on lifestyles, especially related to mating. Several discrete and composite chemical signals have been deciphered in elephants, one of which is a preovulatory female-to-male pheromone, (Z)-7-dodecen-1-yl acetate. This pheromone and other recently described or potential chemical signals are compared to compounds in insect pheromone blends. Such knowledge of the chemical ecology of the Asian elephant has potentially important implications for conservation.

Raubenheimer, E.J., Bosman, M.C., Vorster, R., Noffke, C.E., 1998. Histogenesis of the chequered pattern of ivory of the African elephant (Loxodonta africana). Arch Oral Biol 43, 868-977.
Abstract: This study aimed to propose a hypothesis on the events which lead to the development of the characteristic chequered pattern of elephant ivory. Twenty fragments of ivory and six elephant tusks were obtained through the National Parks Board of South Africa. Polished surfaces were prepared in sagittal and longitudinal planes and the characteristics of the distinctive chequered pattern described. Light- and electron-microscopical techniques and image analyses were employed to determine the morphological basis of the pattern and to describe the spatial distribution, density and morphology of the dentinal tubules. These investigations showed that the distinctive pattern was the result of the sinusoidal, centripetal course followed by dentinal tubules. The apical, slanted part of the sinusoidal curve is the result of the centripetally moving odontoblast, which, during formation of ivory, progresses towards the centre of the tusk on a decreasing circumference. It is suggested that this leads to cell crowding, increased pressure between odontoblasts and subsequent apical movement of their cell bodies, cell degeneration and fusion. Odontoblastic degeneration and fusion probably relieve the pressure between the crowded odontoblasts by reducing their numbers and the remaining odontoblasts now orientate their centripetal course towards the tip of the tusk, thereby forming the anterior-directed part of the sinusoidal path of the tubule. As odontoblasts progress centripetally the diameter of the pulpal cavity decreases further and the processes of apical movement, fusion and degeneration of odontoblasts are repeated. This occurs until the pulpal cavity is obliterated.

Tada, T., Watanabe, Y.H., Matsuoka, A., Ikeda-Saito, M., Imai, K., Ni-hei, Y., Shikama, K., 1998. African elephant myoglobin with an unusual autoxidation behavior: comparison with the H64Q mutant of sperm whale myoglobin. Biochim Biophys Acta 1387, 165-176.
Abstract: Elephant myoglobins both from Asian and African species have a glutamine in place of the usual distal (E7) histidine at position 64. We have isolated native oxymyoglobin directly from the skeletal muscle of African elephant (Loxodonta africana), and examined the autoxidation rate of oxymyoglobin (MbO2) to metmyoglobin (metMb) as a function of pH in 0.1 M buffer at 25 degreesC. As a result, African elephant MbO2 was found to be equally resistant to autoxidation as sperm whale myoglobin. However, the elephant myoglobin exhibited a distinct rate saturation below pH 6. Kinetic analysis of the pH profiles for the autoxidation rate has disclosed that African elephant MbO2 does not show any proton-catalyzed process, such as the one that can play a dominant role in the autoxidation reaction of sperm whale myoglobin by involving the distal histidine as its catalytic residue. Such a greater stability of African elephant MbO2 at low pH could be explained almost completely by the single H64Q mutation of sperm whale myoglobin. In African elephant aqua-metmyoglobin the Soret band was considerably broadened so as to produce another peak in the pentacoordinate 395 nm region. This unique spectral feature was therefore analyzed to show that the myoglobin is in equilibrium between two species, depending upon the presence or absence of a water molecule at the sixth coordinate position.

Welsch, U., Feuerhake, F., van Aarde, R., Buchheim, W., Patton, S., 1998. Histo- and cytophysiology of the lactating mammary gland of the African elephant(Loxodonta africana). Cell Tissue Res 294, 485-501.
Abstract: The lactating mammary gland of the African elephant (Loxodonta africana) has been studied with a panel of morphological techniques focusing on (1) the functional changes during the secretory process, (2) proliferative process [by application of proliferating cell nuclear antigen (PCNA) immunohistochemistry] and apoptotic phenomena [by use of the TUNEL technique] in the individual lobules, and (3) components of milk and milk-fat-globule membrane. In the lactating gland, the lobules are variably differentiated; within a lobule, however, the alveoli are usually similarly differentiated. The morphology of their alveoli suggests a classification of the lobules into types 1-3. Lobules of type 1 are composed of immature tubular alveoli with mitotic figures and numerous PCNA-positive nuclei; advanced type 1 alveoli contain abundant glycogen and specific secretory granules. Lobules of type 2 are further subdivided. In type 2a lobules, the epithelial cells of the alveoli form tall apical protrusions, which in part are occupied by small lipid droplets and which are pinched off in an apocrine fashion. The number of lysosomes varies considerably. Type 2b is the most common type, with striking basal membrane foldings, abundant rough endoplasmic reticulum cisterns, large Golgi apparatus, numerous mitochondria, lipid droplets, and protein vesicles with 30- to 90-nm-wide casein micelles. The lipid droplets are pinched off with minimal amounts of cytoplasm. Type 2c is composed of alveoli with a cuboidal epithelium and few signs of secretory activity. Increasing expression of peanut-agglutinin-binding sites parallels the maturation and differentiation of the glandular cells. Type 3 lobules are marked by numerous TUNEL-positive nuclei and large lipid droplets and are apparently degenerating structures. Cytokeratin (CK) 14 is usually present in the myoepithelial cells; CK 19 and CK 7 mark ductal and immature alveolar epithelia. Milk protein content varies between 2.6% and 6.3%, and casein micelles range from 35 to 90 nm in diameter. The diameter of intra-alveolar milk fat globules ranges from 5 to 25 micrometer and the membranes bear a filamentous surface coat composed of membrane-anchored mucins; gel-electrophoretic analysis of these mucins from different individuals demonstrates the presence of mucin MUC 1, which is expressed with considerable genetic heterogeneity.

Brown, R.E., Butler, J.P., Godleski, J.J., Loring, S.H., 1997. The elephant's respiratory system: adaptations to gravitational stress. Respiratory Physiology 110, 67.
Abstract: Elephants have had to adapt to gravitational stresses imposed on their very large respiratory structures. We describe some unusual features of the elephant's respiratory system and speculate on their functional significance. A distensible network of collagen fibers fills the pleural space, loosely connects lung to chest wall but appears not to constrain lung-chest wall movements. Myriad spaces within the network and its rich supply of capillaries suggest effective local sources and sinks for pleural fluid that may replace the gravity-dependent flows of smaller mammals. The lung is partitioned into approximately equal to 1 cm3 parenchymal units by a system of thick, elastic septa that ramify throughout the lung from origins on the lung's elastic external capsule. Parenchymal units suspended upon the elastic septal system protect dependent alveoli from compression, thereby reducing the usual gravitational gradient of lung expansion. Intra-pulmonary airways are devoid of cartilage, instead they appear to derive resistance to collapse from tethering forces of the attached septa.

Keet, D.F., Grobler, D.G., Raath, J.P., Gouws, J., Carstens, J., Nesbit, J.W., 1997. Ulcerative pododermatitis in free-ranging African elephant (Loxodonta africana) in the Kruger National Park. Onderstepoort Journal of Veterinary Research 64, 25-32.
Abstract: The occurrence of severe lameness in adult African elephant bulls in a shrub Mopane (Colophospermum mopane) ecosystem was investigated. Large ulcers in the soles of at least one front foot were seen in each of the recorded cases. Microscopically, the lesion can be described as a severe, chronic-active, ulcerative, bacterial pododermatitis (complicated by hypersensitivity/septic vasculitis). A variety of bacteria were isolated from these lesions as well as from regional lymph nodes. Streptococcus agalactiae was the most consistent isolate, while Dichelobacter nodosus, the only organism known to be involved with foot disease in domestic ruminants, was isolated from two cases. Contributory factors such as body mass, portal of entry and origin of potential pathogens may have predisposed to the development of the lesions.

Pribe, C., Grossberg, S., Cohen, M.A., 1997. Neural control of interlimb oscillations. II. Biped and quadruped gaits and bifurcations. Biol Cybern 77, 141-152.
Abstract: Behavioral data concerning animal and human gaits and gait transitions are simulated as emergent properties of a central pattern generator (CPG) model. The CPG model is a version of the Ellias-Grossberg oscillator. Its neurons obey Hodgkin-Huxley type equations whose excitatory signals operate on a faster time scale than their inhibitory signals in a recurrent on-center off-surround anatomy. A descending command or GO signal activates the gaits and triggers gait transitions as its amplitude increases. A single model CPG can generate both in-phase and anti-phase oscillations at different GO amplitudes. Phase transitions from either in-phase to anti-phase oscillations or from anti-phase to in-phase oscillations can occur in different parameter ranges, as the GO signal increases. Quadruped vertebrate gaits, including the amble, the walk, all three pairwise gaits (trot, pace, and gallop), and the pronk are simulated using this property. Rapid gait transitions are simulated in the order--walk, trot, pace, and gallop--that occurs in the cat, along with the observed increase in oscillation frequency. Precise control of quadruped gait switching uses GO-dependent modulation of inhibitory interactions, which generates a different functional anatomy at different arousal levels. The primary human gaits (the walk and the run) and elephant gaits (the amble and the walk) are simulated, without modulation, by oscillations with the same phase relationships but different waveform shapes at different GO signal levels, much as the duty cycles of the feet are longer in the walk than in the run. Relevant neural data from spinal cord, globus pallidus, and motor cortex, among other structures, are discussed.

Sarma, K.K., Dutta, B., 1997. Preputial diverticulum in an Asian elephant (Elephas maximus) - a case report. Indian Veterinary Journal 74, 59-60.

Shoshani, J., 1997. Origins and evolution. In: Eltringham, M.A. (Ed.), The illustrated encyclopedia of elephants. Salamander Books Ltd., London, pp. 12-29.

Ullrey, D.E., Crissey, S.D., Hintz, H.F. Elephants: Nutrition and Dietary Husbandry: Nutrition Advisory Group Handbook Fact Sheet 004 September 1997. Nutrition Advisory Group Handbook , 1-18. 1997.
Ref Type: Electronic Citation

Bezuidenhout, A.J., Seegers, C.D., 1996. The osteology of the African elephant (Loxodonta africana): vertebral column, ribs and sternum. Onderstepoort Journal of Veterinary Research 63, 131-147.
Abstract: The vertebral column, sternum and ribs of the African elephant were studied and illustrated. In the cervical series, the vertebrae are characterized by very short (compressed) vertebral bodies and short spinous processes. There are 20-21 thoracic vertebrae that carry ribs, and three lumbar vertebrae. The neural arches of the five sacral vertebrae fuse with each other as well as with the wings of the ilium, while the intervertebral discs do not ossify and the vertebral bodies remain separate. There are 19-21 caudal vertebrae. In the latter, the neural arches of only the first five to six vertebrae fuse dorsally, the vertebral foramens of the other vertebrae as well as the vertebral canal remain open dorsally.The body of the first rib is greatly expanded while that of the last three to four ribs are reduced. The cartilages of the first six ribs articulate with the sternum, the last five to six ribs do not bear costal cartilages and are not attached to the costal arch.The sternum consists of five sternabrae that form three approximately equal, but separate, segments. The first segment is formed by the first sternabra, the second segment is formed by the second to fourth sternabrae and the last segment is formed by the fifth sternabra. The first and second sternabrae articulate with each other by means of a synovial joint, the second to fourth sternabrae are fused to each other and the fourth and fifth sternabrae are loosely attached to each other by connective tissue.

Foged, N.T., Delaisse, J.M., Hou, P., Lou, H., Sato, T., Winding, B., Bonde, M., 1996. Quantification of the collagenolytic activity of isolated osteoclasts by enzyme-linked immunosorbent assay. J Bone Miner Res 11 , 226-237.
Abstract: Difficulties in the geometrical definition and measurement of resorption pits is a major problem for the quantitative analysis of bone resorption by isolated osteoclasts cultured on bone or dentin substrates. In this study we developed an enzyme-linked immunosorbent assay (ELISA) for quantification of bone resorption in vitro, which specifically quantifies type I collagen fragments released into the culture medium by the resorptive action of bone cells cultured on slices of bone or dentin. A consistently high correlation between the formation of resorption pits and the release of antigenic collagen fragments was observed for isolated rabbit osteoclasts seeded at various densities and cultured for various periods on bovine, elephant, and human substrates. In a further support of the osteoclastic nature of the collagenolytic effects, a high consistency between pit formation and collagenolysis was also observed when the rabbit bone cells were cultured in the presence of very differently acting but typical inhibitors of pit formation, i.e., the carbonic anhydrase inhibitor acetazolamide, the cysteine proteinase inhibitor epoxysuccinyl-L-leucylamido-(4-guanodino)butane (E-64), the phosphatidyl-inositol 3-kinase inhibitor wortmannin, and the bisphosphonate ibandronate (BM 21.0955). In conclusion, the ELISA represents a simple, precise, and objective way to dynamically monitor bone resorption in vitro through quantification of the collagenolytic activity of isolated osteoclasts.

Kaufman, M.H., 1996. Observations on Barclay's elephant. J R Coll Surg Edinb 41, 75-81.
Abstract: This account attempts to trace the fate of the skeleton of an elephant that was gifted by George Ballingall to Dr John Barclay, one of the most important teachers of Anatomy in Edinburgh during the early 19th century. In his will, elephant, to the Royal College of Surgeons of Edinburgh on two conditions, that a hall should be built to house the collection, and that it should be associated with his name in perpetuity. In the 1830s, the comparative collection, but particularly the skeleton of the elephant, was the pride of the College. Unfortunately, interest in the comparative material rapidly diminished, and, due to constraints on space, while the elephant's skull was retained the rest of the skeleton was disposed of. An unpublished poem written at the time of the Burke trial, in 1829, testifies to the fact that Barclay's elephant was closely associated in the minds of the public with the activities of Dr Robert Knox, the then Conservator of the College museum.

Lewandowski, K., Busch, T., Lewandowski, M., Keske, U., Gerlach, H., Falke, K.J., 1996. Evidence of nitric oxide in the exhaled gas of Asian elephants (Elephas maximus). Respiratory Physiology 106, 91-98.
Abstract: Nitric oxide (NO) produced in the respiratory tract is released into the respiratory gases of humans, rabbits, guinea-pigs, and rats. We analysed the NO concentrations in the exhaled gas of four awake Asian elephants. Two methods were employed: (1) exhaled gas was sampled from the elephants' trunks with a 1 L syringe and analysed for NO concentrations by chemiluminescence; (2) respiratory gas was continuously aspirated via a thin plastic tube positioned within the trunk and on-line analysed for NO concentrations by chemiluminescence. Syringe sampling (n = 4), when corrected for dilution by ambient air using linear regression analysis, revealed a mean NO concentration of 31 parts per billion (ppb); highest exhalatory concentrations measured during continuous suctioning were 27 and 28 ppb (n = 2). The exhaled NO concentrations in elephants are similar to those found in humans measured with a comparable technique. This supports the hypothesis that a size-independent 'normal value' of endogenous NO is provided in the airways which may contribute to regulation of pulmonary ventilation and perfusion by autoinhalation in some mammals.

Soltysiak, Z., 1996. Age-related changes in the brain of an Indian elephant. Zycie-Weterynaryjne 71, 309-311.

Van Aswegen, G., Van Noorden, S., Kotze, S.H., de Vos, V., Schoeman, J.H., 1996. The intestine and endocrine pancreas of the African elephant: a histological, immunocytochemical and immunofluorescence study. Onderstepoort Journal of Veterinary Research 63, 335-340.
Abstract: Histological, immunocytochemical and immunofluorescence methods were employed to study the intestine and endocrine pancreas of the elephant. The histological findings were in line with those in monogastric animals. In the mucosa of intestine, endocrine cells were immunoreactive to somatostatin, gastrin, CCK, GIP, secretin, motilin, glucagon and NPY. Nerve cells immunoreactive to somatostatin, substance P, VIP, PHI, NPY, bombesin and CGRP were detected. No immunoreactivity to neurotensin was observed. Islets of the pancreas had insulin cells in their cores and glucagon and somatostatin cells in their mantles. The antisera employed failed to demonstrate PP cells in the pancreas, but NPY-immunoreactive cells were present.

Bisig, D.A., Di Iorio, E.E., Diederichs, K., 1995. Crystal structure of Asian elephant (Elephas maximus) cyanometmyoglobin at 178-A resolution. Phe29(B10) accounts for its unusual ligand binding properties. Journal of Biological Chemistry 270, 20752-20754.
Abstract: The crystal structure of Asian elephant cyano-metmyoglobin which has a glutamine instead of the usual distal site histidine has been determined to high resolution. In addition to this replacement, the substitution of a conserved leucine residue in position 29(B10) at the distal side by a phenylalanine was unambiguously identified based on the available electron density. The suspicion, that there were errors in the original sequence which has caused some confusion, is thus confirmed. Comparison with other myoglobin structures in various ligated forms reveals an essentially unchanged tertiary structure in elephant myoglobin despite the two amino acid substitutions in the heme pocket. Our current structural model shows that the N epsilon 2 atom of Gln64(E7) has moved with respect to the corresponding nitrogen position of His64(E7) in the CO complex of sperm whale myoglobin. The newly assigned residue Phe29(B10) penetrates into the distal side of the heme pocket approaching the ligand within van der Waals distance and causing a much more crowded heme pocket compared to other myoglobins. Kinetic properties of Asian elephant myoglobin, wild type, and recombinant sperm whale myoglobins are discussed in relation to the structural consequences of the two amino acid substitutions H64Q and L29F.

Coetzee, H.L., Kotze, S.H., Lourens, N., 1995. Characterization of mucus glycoproteins in the intestinal mucosa of the African elephant (Loxodonta africana) following lectin histochemistry. Onderstepoort Journal of Veterinary Research 62, 187-192.
Abstract: The glycoproteins of the small intestines, caecum and colon of three adult elephants and one recently weaned elephant calf were examined by means of lectin histochemistry. Tissue sections were histochemically stained with peroxidase-labelled concanavalin A (Con A), asparagus-pea (TPA), peanut (PNA) and wheat-germ (WGA) lectins. Con A and TPA showed no binding activity in the intestinal tract of the adult elephants or the duodenum and ileum of the elephant calf, but did show a small amount of binding activity in the caecum and colon of the calf. WGA bound very intensely throughout the intestinal tracts of the adults and of the calf--especially with the goblet cells located in the crypts of Lieberkuhn and the glands of Brunner--decreasing in intensity towards the luminal surface of the intestinal tract. PNA stained the glands of Brunner of the duodenum faintly and the goblet cells of the ileum moderately, with no staining of the caecum and faint staining of the colon. These results show the distribution of Con A-, WGA-, PNA- and TPA-binding sites, and the changes that take place in the type of glycoprotein secreted after a change in the diet of the animal.

Endo, H., Yamada, T.K., Suzuki, N., Suwa, G., Uetsuka, K., Hashimoto, O., Kurohmaru, M., Hayashi, Y., 1995. Ultrastructure of cardiac myocyte in the Asian elephant (Elephas maximus). Journal of Veterinary Medical Science 57, 1035-1039.
Abstract: Cardiac myocytes of an Asian elephant (Elephas maximus) were observed by transmission electron microscopy. Typical ultrastructural features of cardiac myocytes are exhibited in the musculature of both the left and right atria, and left ventricle of the heart. Myofibrils, mitochondria, T-system and sarcoplasmic reticulum are well-developed within the cytoplasm. Many mitochondria are characteristically concentrated is some myocytes. Cardiac musculature is also distributed in the root of the caudal vena cava. Many atrial granules are detected not only in atrial myocytes, but also in the myocytes of the caudal vena cava. Atrial natriuretic polypeptide may be secreted from the caval venous wall in the elephant.

Kramer, B., Hattingh, J., 1995. The neuromuscular junction in the African elephant Loxodonta africana and African buffalo Syncerus caffer. South African Journal of Wildlife Research 25, p14, 3p, 2bw.
Abstract: Differences in the physiological response to the drug succinyldicholine occur between the African elephant Loxodonta africana and African buffalo Syncerus caffer, irrespective of the route of administration of the drug. The response in elephants has suggested the presence of unique acetylcholine receptors in their respiratory muscles. In this paper the first observations of the neuromuscular junction in the African elephant and African buffalo are reported. While the basic structure of the junction was found to be typically mammalian in both species, differences were found in the morphology of the postjunctional area where these receptors reside. Elucidation of the structure and function of this junction in these animals is important in the selection of drugs that act as neuromuscular blockers.

Langman, V.A., Roberts, T.J., Black, J., Maloiy, G.M.O., Heglund, N.C., Weber, J.M., Kram, R., Taylor, C.R., 1995. Moving cheaply: energetics of walking in the African elephant. J Exp Biol 198 (Pt 3 ), 629-632.
Abstract: Large animals have a much better fuel economy than small ones, both when they rest and when they run. At rest, each gram of tissue of the largest land animal, the African elephant, consumes metabolic energy at 1/20 the rate of a mouse; using existing allometric relationships, we calculate that it should be able to carry 1 g of its tissue (or a load) for 1 km at 1/40 the cost for a mouse. These relationships between energetics and size are so consistent that they have been characterized as biological laws. The elephant has massive legs and lumbers along awkwardly, suggesting that it might expend more energy to move about than other animals. We find, however, that its energetic cost of locomotion is predicted remarkably well by the allometric relationships and is the lowest recorded for any living land animal.

Maluf, N.S.R., 1995. Kidney of elephants. Anatomical Record 242, 491-514.
Abstract: BACKGROUND: Elephants are an important and isolated order. Their kidneys need substantial investigation and hitherto have not been portrayed even by a pyelogram. METHODS: Pyelograms and injection of vessels with colored acrylic emulsions were done initially. Dissection was under fiberoptics using a dissecting microscope with frequent measurements. Special areas were cut for microscopy (light and electron) and photography. Glomerular counts were done by macerating weighted pieces of cortex and later finding the cortical fraction of the renal parenchyma. RESULTS: The elephant kidney is devoid of dorsoventral symmetry. It is composed of 8 +/- 2 lobes separated by fine interlobar septa. There is no reduction of lobes with maturity. The pelvis bifurcates at the sinus into primary branches or infundibula which dispatch a secondary branch or infundibulum into every lobe. Interlobar arteries and veins, nerves, fat, and connective tissue generally accompany every secondary infundibulum into its lobe. A major branch of the renal artery may perforate the renal capsule and course to the cortico-medullary (C-M) border independently of the secondary infundibulum to that lobe. The number of glomeruli per kidney is approximately 15 x 10(6). In adults the glomerular mass is 4.9 +/- 0.5% of the renal parenchyma and 6.7 +/- 0.3% of the cortex. Areae cribrosae occur generally at low papillae. They are the outlets of numerous terminal collecting ducts which may be accompanied by a tubus maximus (T.M.) A T.M. of diameter 1.6 mm and length 10 mm may act as the only substitute for an area cribrosa. Wide anastomoses between the two main renal veins occur within the renal sinus. Intralobar arteries and veins often course right through the outer medulla to and from, respectively, the C-M border. CONCLUSIONS: Anatomically, an elephant's kidneys appear to be able to concentrate urine only moderately. Their kidneys tend to resemble those of the manatee but not of the dugong.

Nummela, S., 1995. Scaling of the mammalian middle ear. Hear Res 85, 18-30.
Abstract: This study considers the general question how animal size limits the size and information receiving capacity of sense organs. To clarify this in the case of the mammalian middle ear, I studied 63 mammalian species, ranging from a small bat to the Indian elephant. I determined the skull mass and the masses of the ossicles malleus, incus and stapes (M, I and S), and measured the tympanic membrane area, A1. The ossicular mass (in mg) is generally negatively allometric to skull mass (in g), the regression equation for the whole material (excluding true seals) being y = 1.373 x(0.513). However, for very small mammals the allometry approaches isometry. Within a group of large mammals no distinct allometry can be discerned. The true seals (Phocidae) are exceptional by having massive ossicles. The size relations within the middle ear are generally rather constant. However, the I/M relation is slightly positively allometric, y = 0.554 x(1.162). Two particularly isometric relations were found; the S/(M + I) relation for the ossicles characterized by the regression equation y = 0.054 x(0.993), and the relation between a two-dimensional measure of the ossicles and the tympanic membrane ares, (M + I)2/3 /A1. As in isometric ears the sound energy collected by the tympanic membrane is linearly related to its area, the latter isometry suggests that, regardless of animal size, a given ossicular cross-sectional area is exposed to a similar sound-induced stress. Possible morphological middle ear adaptations to particular acoustic environments are discussed.

Raubenheimer, E.J., van Heerden, W.F., van Niekerk, P.J., de Vos, V., Turner, M.J., 1995. Morphology of the deciduous tusk (tush) of the African elephant (Loxodonta africana). Arch Oral Biol 40, 571-576.
Abstract: The tusk of the African elephant is preceded by a deciduous tooth generally known as the tush. Tushes from nine elephant fetuses and six calves younger than 1 year were exposed by dissection and described morphologically. All tushes consisted of a crown, root and pulpal cavity, the formation of which is completed soon after birth. They reached a maximum length of 5 cm, appeared not to erupt through the skin and were pushed aside and resorbed during enlargement of the distally located primordium of the tusk. Dental enamel, which covered the crown, could easily be removed and consisted of rods with an interwoven arrangement; the dentine-enamel junction was flat. Cellular cementum extended for variable distances over the crown and the dentine was tubular in nature. Although the tush apparently has no function, it provides the anlage and orientation for the development of its permanent successor.

Van-der-Merwe, N.J., Bezuidenhout, A.J., Seegers, C.D., 1995. The skull and mandible of the African elephant (Loxodonta africana). Onderstepoort Journal of Veterinary Research 62, 245-260.
Abstract: In the present study the bones of the skull, excluding the hyoid apparatus, are described. All the bones are aerated by sinuses. In the occipital bone the squamous part is aerated from the sinus of the parietal bone, the lateral part is aerated from the tympanic bulla and the basal part from the sinus of the basisphenoid bone. Condylar foramens and hypoglossal canals are absent. A small interparietal bone is present at birth. At an early age it fuses with the surrounding cranial bones. The squamous part of the temporal bone lies sagittally in young animals, but moves progressively to a transverse plane as the animals age. A foramen lacerum is represented by jugular and oval foramens and the carotid canal. The body of the basisphenoid bone is excavated by the massive maxillary tuberosity. The latter extends to the oval foramen and contains the developing molar teeth. The ethmoturbinate, nasal and lacrimal bones are exceptionally small. In old bulls the palatine process of the incisive bones and their sinuses are gradually displaced by the palatine process of the maxillae.

Kotze, S.H., Coetzee, H.L., 1994. A histocytochemical study of mucus glycoproteins or mucins in the intestinal tract of the African elephant (Loxodonta africana). Onderstepoort Journal of Veterinary Research 61, 177-181.
Abstract: The distribution of neutral mucins, sialomucins and sulphomucins was determined histochemically in the duodenum, jejunum, ileum and colon of the African elephant (Loxodonta africana). The techniques used were periodic acid-Schiff (PAS), alcian blue/periodic acid-Schiff (AB-PAS), high-iron-diamine/alcian blue (HID-AB), alcian blue at varying pH solutions and alcian blue at high temperature after methylation and saponification. Acid mucins appear to dominate neutral mucins, the latter decreasing toward the large intestine. Sulphomucins and sialomucins occurred in almost equal amounts throughout the intestinal tract, with a slight decrease of sialomucins toward the colon.

Ringo, J.L., Doty, R.W., Demeter, S., Simard, P.Y., 1994. Time is of the essence: a conjecture that hemispheric specialization arises from interhemispheric conduction delay. Cereb Cortex 4, 331-343.
Abstract: Tomasch (1954) and Aboitiz et al. (1992) found the majority of the fibers of the human corpus callosum are under 1 micron in diameter. Electron microscopic studies of Swadlow et al. (1980) and the detailed study of LaMantia and Rakic (1990a) on macaques show the average size of the myelinated callosal axons also to be less than 1 micron. In man, the average-sized myelinated fiber interconnecting the temporal lobes would have a one-way, interhemispheric delay of over 25 msec. Thus, finely detailed, time-critical neuronal computations (i.e., tasks that strain the capacity of the callosum and hence could not be handled by just the larger fibers) would be performed more quickly via shorter and faster intrahemispheric circuits. While one transit across the commissural system might yield tolerable delays, multiple passes as in a system involving "setting" would seem prohibitively slow. We suggest that these temporal limits will be avoided if the neural apparatus necessary to perform each high-resolution, time-critical task is gathered in one hemisphere. If the, presumably overlapping, neural assemblies needed to handle overlapping tasks are clustered together, this would lead to hemispheric specialization. The prediction follows that the large brains of mammals such as elephants and cetaceans will also manifest a high degree of hemispheric specialization.

Shoshani, J., 1994. Skeletal and other basic anatomical features of elephants. In: Shoshani, J., Tassy, P. (Eds.), The proboscidea: evolution and paleoecology of elephants and their relatives. Oxford University Press, Oxford, pp. 9-20.

Smuts, M.M.S., Bezuidenhout, A.J., 1994. Osteology of the pelvic limb of the African elephant (Loxodonta africana). Onderstepoort Journal of Veterinary Research 61, 51-66.
Abstract: The pelvic girdle was characterized by large, transversely-placed ilial wings. The femur was the longest bone of the skeleton and its fovea capitis was situated caudomedially between the epiphyseal line and the articular surface of the femoral head. A wedge-shaped patella articulated with the femoral trochlea. The bones of the crus were approximately half as long as the femur and consisted of the sturdy tibia and slender fibula. The condyles of the tibia were concave and the femoro-tibial joint was congruent with rudimentary menisci. The tarsus consisted of seven bones which were arranged in three rows. There were five metatarsal bones. Only four digits were present, the third and fourth consisted of three phalanges each while the second and fourth digits were smaller and consisted of two phalanges each. The first digit was represented by one proximal sesamoid bone only. A large, cartilagenous rod or pre-hallux was attached to the first tarsal and metatarsal bones. Proximal sesamoid bones were present on the plantar aspect of the trochleae of metatarsal bones 1-V. The pes was found to be digitigrade and the digits rested on a thick pad of elastic connective tissue and fat.

Van-Aswegen, G., Schoeman, J.H., De-Vos, V., Van-Noorden, S., 1994. The oesophagus and stomach of the African elephant: a histological, immunocytochemical and immunofluorescence study. Onderstepoort Journal of Veterinary Research 61, 223-229.
Abstract: Histological, immunocytochemical and immunofluorescence methods were employed to study the oesophagus and stomach of the elephant. The histological findings were similar to those in monogastric species like pigs and humans. In the mucosa of the stomach, endocrine cells were immunoreactive to gastrin, somatostatin, chromogranin A and serotonin. Nerve cells immunoreactive to somatostatin, bombesin, VIP, PHI and CGRP were detected in the submucosal and myenteric plexus of the stomach. In the stomach, the absence of glucagon cells and the presence of endocrine cells immunoreactive to PYY, are in contrast to the situation in other mammals and need further investigation. Small gastric ulcers were observed in some of the specimens.

Bechert, U.S., 1993. Morphology and physiology. Animal Keepers' Forum 20, 111-115.

Cupane, A., Leone, M., Vitrano, E., Cordone, L., Hiltpold, U.R., Winterhalter, K.H., Yu, W., DiIorio, E.E., 1993. Structure-dynamics-function relationships in Asian elephant (Elephas maximus)myoglobin. An optical spectroscopy and flash photolysis study on functionally important motions. Biophys J 65, 2461-2472.
Abstract: In this work we report the thermal behavior (10-300 K) of the Soret band lineshape of deoxy and carbonmonoxy derivatives of Asian elephant (Elephas maximus) and horse myoglobins together with their carbon monoxide recombination kinetics after flash photolysis; the results are compared to analogous data relative to sperm whale myoglobin. The Soret band profile is modeled as a Voigt function that accounts for the coupling with high and low frequency vibrational modes, while inhomogeneous broadening is taken into account with suitable distributions of purely electronic transition frequencies. This analysis makes it possible to isolate the various contributions to the overall lineshape that; in turn, give information on structural and dynamic properties of the systems studied. The optical spectroscopy data point out sizable differences between elephant myoglobin on one hand and horse and sperm whale myoglobins on the other. These differences, more pronounced in deoxy derivatives, involve both the structure and dynamics of the heme pocket; in particular, elephant myoglobin appears to be characterized by larger anharmonic contributions to soft modes than the other two proteins. Flash photolysis data are analyzed as sums of kinetic processes with temperature-dependent fractional amplitudes, characterized by discrete pre-exponentials and either discrete or distributed activation enthalpies. In the whole temperature range investigated the behavior of elephant myoglobin appears to be more complex than that of horse and sperm whale myoglobins, which is in agreement with the increased anharmonic contributions to soft modes found in the former protein. Thus, to satisfactorily fit the time courses for CO recombination to elephant myoglobin five distinct processes are needed, only one of which is populated over the whole temperature range investigated. The remarkable convergence and complementarity between optical spectroscopy and flash photolysis data confirms the utility of combining these two experimental techniques in order to gain new and deeper insights into the functional relevance of protein fluctuations.

Kern, T.J., Murphy, C.J., Howland, H.C. Physiological optics and ocular anatomy of the Asian elephant (Elephas maximus). Proceedings of the American Association of Zoo Veterinarians. 355. 1993.
Ref Type: Conference Proceeding

Rasmussen, L.E.L., Johnson, E.W., Jafek, B.W. Preliminary observations on the morphology of the vomeronasal organ of a newborn Asian elephant. Chemical Senses 18, 618. 1993.
Ref Type: Abstract
Abstract: Abstract. Full-text. Adult Asian elephants have an apparently typical mammalian vomeronasal organ (VNO). Presumably, flehmen responses aid in the presentation of bioactive molecules to vomeronasal neuroreceptors. Young Asian elephants do not exhibit flehmen responses until 6-17 weeks after birth. Histological studies of VNO in newborn elephants have not been available. Recently, at the light microscopic level, we have observed a structure that in gross appearance is similar to the VNO of other mammals; there is a lumen surrounded by a convex and a concave epithelial border, those borders join at both ends. Based on previous studies, we presume that the concave border would be the neuroepithelium with the receptor cells. The epithelia of both surfaces are pseudostratified. Within these epithelia are cells with different nuclear morphologies. Some of the nuclei are euchromatic and oval. Other appear heterochromatic. Round basal cells are also apparent. At the surfaces of the epithelia some ciliated cells can be seen. To our knowledge, ciliated cells have been identified in the VNO neuroepithelium of only one other mammalian species. To further document the cell types found in the newborn elephant VNO and to attempt to identify receptor cells, we will do electron microscopy on representative regions.

Smuts, M.M.S., Bezuidenhout, A.J., 1993. Osteology of the thoracic limb of the African elephant (Loxodonta africana). Onderstepoort Journal of Veterinary Research 60, 1-14.
Abstract: The forelimb bones of 8 elephants (7 adults, 1 juvenile) were studied. In addition, the bones of the digits were dissected and studied in situ in a mature specimen. The scapula, humerus and bones of the antebrachium (particularly the ulna) are massive in comparison to the short, relatively small bones of the manus. There are 8 carpal bones, 5 metacarpal bones and 5 digits. Digits 2-4 consist of 3 phalanges each. The 5th digit consists of 2 phalanges, while the 1st is represented by a single phalanx which is tusk-like and pointed. The distal phalanges of digits 2-4 are very small and do not articulate with the middle phalanges. The proximal sesamoids are well developed and are present on the palmar aspect of all 5 metacarpophalangeal joints. All the bones are illustrated from at least 2 aspects.

Vyas, K., Rajarathnam, K., Yu, L.P., Emerson, S.D., La Mar, G.N., Krishnamoorthi, R., Mizukami, H., 1993. 1H NMR investigation of the heme cavity of elephant (E7 Gln)met-cyano-myoglobin. Evidence for a B-helix phenylalanine interaction with bound
ligand. J Biol Chem 268, 14826-14835.
Abstract: A combination of one- and two-dimensional NMR experiments has been used to identify and spatially locate the heme pocket residues in the paramagnetic, low spin, met-cyano complex of elephant myoglobin. In addition to assigning resonances of the conserved residues, we have also assigned Gln64 (E7) and an aromatic ring designated PheA whose side chain is inserted into the heme pocket, as found earlier for elephant carbonmonoxy-myoglobin and oxy-myoglobin (Yu, L. P., La Mar, G. N., and Mizukami, H. (1990) Biochemistry 29, 2578-2585). The assigned conserved proximal side residues (Leu89(F4), Ala90(F5), His93(F8), His97(FG3), Ile99(FG5), Leu104(G5), Phe138(H15), and Tyr146(H23)) and conserved distal side residues (Phe43(CD1), Thr67(E10), Val68(E11), and Ala71(E14)) in elephant met-cyano-myoglobin are found to have orientations similar to those in sperm whale met-cyano-myoglobin. The observed dipolar connectivities and dipolar shift pattern for the substituted Gln64(E7) place the Gln in the heme pocket oriented toward the iron, as found for His64(E7). The conserved structural elements demand that the inserted PheA originate from the B-helix (i.e. Phe27 or Phe33). Dipolar contacts between the inserted PheA and the conserved residues Phe43(CD1), Val68(E11), Ile107(G8), and Gln64(E7), place PheA in the position occupied by the B10 residue in sperm whale myoglobin (Mb), with the larger size of the PheA side chain as compared to the replaced Leu being accommodated by the vacancy that occurs in sperm whale Mb. The paramagnetic induced relaxation places PheA in van der Waals contact with the bound ligand. Hence we conclude that the B10 position of elephant Mb is occupied by a Phe, and this substitution relative to sperm whale Mb is responsible for the low autoxidation rate and low reduction potential of elephant Mb. A reduced autoxidation rate has been reported for a sperm whale synthetic point mutant Leu29(B10) --> Phe (Carver, T. E., Brantley, R. E., Jr., Singleton, E. W., Arduini, R. M., Quillin, M. L., Phillips, G. N., and Olson, J. S. (1992) J. Biol. Chem. 267, 14443-14450). The published sequence of elephant Mb places B-helix Phe residues at position 27(B8) and 33(B14), but a Phe at neither of these positions can account for the observed NMR properties. Since a large proportion of the substitutions in elephant relative to sperm whale Mb, and some of the least conservative, occur in the B-helix, neither a structurally perturbed B-helix nor an error in the sequence can be discounted.

Chungath, J.J., Paily, L., Ommer, P.A., 1992. Anatomy of the vertebral column of the Indian elephant (Elephas maximus). In: Silas, E.G., Nair, M.K., Nirmalan, G. (Eds.), The Asian Elephant: Ecology, Biology, Diseases, Conservation and Management (Proceedings of the National Symposium on the Asian Elephant held at the Kerala Agricultural University, Trichur, India, January 1989). Kerala Agricultural University, Trichur, India, pp. 43-45.

Harshan, K.R., Chungath, J.J., Paily, L., Ommer, P.A., 1992. Histology of the temporal gland of the Indian elephant (Elephas maximus). In: Silas, E.G., Nair, M.K., Nirmalan, G. (Eds.), The Asian Elephant: Ecology, Biology, Diseases, Conservation and Management (Proceedings of the National Symposium on the Asian Elephant held at the Kerala Agricultural University, Trichur, India, January 1989). Kerala Agricultural University, Trichur, India, pp. 46-48.

Hattingh, J., Petty, D., 1992. Comparative physiological responses to stressors in animals. Comparative Biochemistry and Physiology A-Comparative-Physiology 101, 113-116.
Abstract: The species-specific experimental response to stressors (SSERTS) analysis was applied to a number of species under varied short and long term conditions. The measure provides quantitative data relating to the physiological responses of animals when exposed to stressors and results are presented comparing these for different methods of immobilization, euthanasia, etc. at intra- and inter-species level. It is suggested that the SSERTS measure is of greater value for measuring the responses of animals to stressors than is the measurement of the concentration of single blood variables.

Murphy, C.J., Kern, T.J., Howland, H.C., 1992. Refractive state, corneal curvature, accommodative range and ocular anatomy of the Asian elephant (Elephas maximus). Vision Res 32, 2013-2021.
Abstract: The resting refractive state of six mature, female, Asian elephants (Elephas maximus) was determined using streak retinoscopy and neutralizing video retinoscopy. The amplitude of accommodation was also measured by neutralizing video retinoscopy of two animals and the corneal curvatures of three animals was measured by photokeratoscopy. The net spherical refraction was found to be +0.23 D. No difference was observed between cyclopleged and non-cyclopleged eyes (data from three animals), nor was there any difference between right and left eyes. Nine of the twelve eyes refracted had > or = 0.5 D astigmatism. The mean corneal power, as measured by photokeratometry was 21.3 D (SD = 1.8 D). There was a tendency towards with-the-rule corneal astigmatism in our sample (mean value: 1.2 D), though it did not reach statistical significance (P = 0.06). Two elephants were examined using neutralizing video photoretinoscopy. They were able to accommodate through 3 D. Three fixed eyes from three different elephants were obtained for gross and microscopic examination. The mean axial length of the eye was 38.75 mm and the lens had an axial diameter of approx. 10 mm. The posterior sclera was thick (8.0-8.5 mm). Histologically, the cornea was comprised of five distinct layers. A thin, meridionally oriented smooth ciliary muscle was identified. Individual muscle fibers were also observed associated with the posterior trabeculae of the uveal meshwork.

Ommer, P.A., Harshan, K.R., Chungath, J.J., Paily, L., 1992. Histology of the adrenal gland of the Indian elephant (Elephas maximus). In: Silas, E.G., Nair, M.K., Nirmalan, G. (Eds.), The Asian Elephant: Ecology, Biology, Diseases, Conservation and Management (Proceedings of the National Symposium on the Asian Elephant held at the Kerala Agricultural University, Trichur, India, January 1989). Kerala Agricultural University, Trichur, India, pp. 49-50.

Phillips, P.K., Heath, J.E., 1992. Heat exchange by the pinnae of the African elephant (Loxodonta africana). Comparative Biochemistry and Physiology [A] 101, 693-699.
Abstract: 1. Surface temperatures of the pinnae of four female African elephants were measured at ambient temperatures between 14 and 32 degrees C using infrared thermography. Instantaneous heat losses calculated using those values ranged from 10.67 to 76.2 W under the observed conditions. 2. Using a value of 17 kcal/kg/day, those heat losses account for 0.65-4.64% of the animals' standard metabolic rates, considering one side of one ear only. 3. A model of heat flow across a flat vertical plate was constructed and compared to the actual values. Up to 100% of an African elephant's heat loss needs can be met by movement of its pinnae and by vasodilation. 4. Thermography indicates that the temperature distribution pattern across the pinna changes with ambient temperature and that areas of specialized motor control exist

Shoshani, J., 1992. Anatomy and physiology. In: Shoshani, J. (Ed.), Elephants. Majestic creatures of the wild. Rodale Press, Emmaus, Pennsylvania, pp. 66-81.

Teunissen, M.J., de Kort, G.V., Op den Camp, H.J., Huis in 't Veld, J.H., 1992. Production of cellulolytic and xylanolytic enzymes during growth of the anaerobic fungus Piromyces sp. on different substrates. J Gen Microbiol 138 (Pt 8), 1657-1664.
Abstract: Piromyces sp. strain E2, an anaerobic fungus isolated from an Indian elephant (hindgut fermenter) was tested for its ability to ferment a range of substrates. The fungus was able to use bagasse, cellobiose, cellulose, fructose, glucose, lactose, mannose, starch, wheat bran, wheat straw, xylan and xylose. Formate and acetate were the main fermentation products after growth on these substrates. The amount of carbon found in the fermentation products of cultures, in which substrate digestion was complete averaged 88.5 mM, or 59% of the carbon offered as substrate. No growth was observed on other substrates tested. Lactose, starch, cellobiose and filter paper cellulose were good inducers of cellulolytic and xylanolytic enzymes. Cellulolytic and xylanolytic enzymes were produced constitutively by Piromyces strain E2, although enzyme activities were generally lower after growth on glucose and other soluble sugars. Complex substrates (bagasse, wheat bran, and wheat straw) were good inducers for xylanolytic enzymes but not for cellulolytic enzymes. The extracellular protein banding pattern after SDS-PAGE was therefore only slightly affected by the growth substrate. Identical beta-glucosidase and endoglucanase activity patterns were found after growth on different substrates. This indicated that differences in enzyme activities were not the result of secretion of different sets of isoenzymes although it remains possible that the relative amount of each isoenzyme produced is influenced by the growth substrate.

Wiesner, H., 1992. Occurrence of Arcus scleralis in elephants. Zoologische Garten 62, 287-293.

Cheng, H.C., Yamashiro, D., 1991. Synthesis and receptor binding activity of elephant beta- endorphin, a beta-endorphin homolog with highly potent analgesic activity. International Journal of Peptide and Protein Research 38, 66-69.
Abstract: Elephant beta-endorphin and its analog, elephant beta- endorphin(6-31) were synthesized by standard solid phase method. Receptor binding activity showed that elephant beta-endorphin was five to six times more potent than human beta-endorphin in its ability to bind to opiate receptors on rat brain membrane. In a previous study (Wong, C.-L., Wai, M.-K., Cheng, H.-C., Chung, D. & Yamashiro, D (1990) Clinical and Experimental Pharmacology and Physiology 16, 33-37), tail flick test for intracerebroventricularly administered beta-endorphin showed that the antinociceptive potency of elephant beta-endorphin was seven to eight times higher than that of human beta-endorphin in mice. Results from both studies suggest that elephant beta-endorphin was a much more potent antinociceptive agent than human beta- endorphin in tail flick test and its higher analgesic activity might be due to its higher affinity for opiate receptors in the brain.

John, M.C., Suramanian, R., 1991. The elephant. Zoos' Print Journal 1-4.

Kozaki, M., Oura, R., Sekine, J., 1991. Studies on digestion physiology of herbivorous feral animals. 2. The comparison of intake of total digestible nutrients among diverse sizes of ruminant and monogastric animals. Journal of the Faculty of Agriculture -Tottori-University 27, 61-68.
Abstract: Digestion trials were carried out on elephant, zebra, giraffe, eland, blackbuck, zebu cattle, Japanese serow, sika deer, muntjac and Japanese Black steer during 3 different seasons of the year. Digestibility of organic matter was about 0.6 for all animals except elephant, muntjac and blackbuck. Crude protein (CP) digestibility correlated (P<0.01) with CP concentration in the feed ration. Acid detergent fibre digestibility ranged from 0.3 to 0.4 in ruminants compared with 0.1 to 0.2 in monogastric animals. No seasonal effects on digesta were observed.

Kramer, B., Teixeira, M., Hattingh, J., 1991. The histology of the adrenal gland of the African elephant, Loxodonta africana. South African Journal of Zoology 26, 193-198.
Abstract: The histology, particularly the ultrastructural cytology, of the adrenal gland of the African elephant, Loxodonta africana, is virtually unknown. Tissue from 14 adult male and female elephants was processed for light and transmission electron microscopy. The gland is surrounded by a thick capsule composed of an outer layer of dense connective tissue and an inner layer in which smooth muscle fibres predominate. Below the layer of smooth muscle, a continuous layer of relatively undifferentiated "capsular" cells occur. Where the capsular cells abut on the zona glomerulosa, they appear to be differentiating into glomerulosa cells, as small lipid droplets are present in their cytoplasm. The cortex is divided into three zones as is found in the adrenal glands of other mammals. Large amounts of collagenous and reticular tissue support the secretory cells, which have a marked lipid content. With electron microscopy, the cortical cells show features typical of steroid-producing cells. The medulla is characterized by an outer region of pale-staining chromaffin-positive (adrenaline) cells and an inner region of intensely staining chromaffin-positive (noradrenaline) cells. The latter cells contain granules of different sizes and structure.

Kroll, W. The straight-tusked elephant of Crumstadt. A contribution to the osteology of Elephas (Palaeoloxodon) antiquus Falconer & Cautley (1847). Der Waldelefant von Crumstadt. Ein Beitrag zur Osteologie des Waldelefanten, Elephas (Palaeoloxodon) antiquus Falconer & Cautley (1847). 1-106. 1991. Munchen, Germany, Tierarztliche Fakultat, Ludwig-Maximilians-Universitat.
Ref Type: Thesis/Dissertation

Ratnasooriya, W.D., Fernando, S.B.U., Manatunga, A.N.V.R., 1991. Presence of an arcus senilis-like structure in the eyes of Sri Lankan elephants (Elephas maximus maximus). Med. Sci. Res. 19, 715-716.

Teunissen, M.J., Smits, A.A.M., Op den Camp, H.J.M., Huis in't Veld, J.H.J., Vogels, G.D., 1991. Fermentation of cellulose and production of cellulolytic and xylanolytic enzymes by anaerobic fungi from ruminant and non- ruminant herbivores. Arch. Microbiol. 156, 290-296.
Abstract: Four anaerobic fungi were grown on filter paper cellulose and monitored over a 7-8 days period for substrate utilization, fermentation products, and secretion of cellulolytic and xylanolytic enzymes. Two of the fungi (N1 and N2) were Neocallimastix species isolated from a ruminant (sheep) and the other two fungi were Piromyces species (E2 and R1) isolated from an Indian Elephant and an Indian Rhinoceros, respectively. The tested anaerobic fungi degraded the filter paper cellulose almost completely and estimated cellulose digestion rates were 0.25, 0.13, 0.21 and 0.18 g.l-1.h-1 for strains E2, N1, N2, R1, respectively. All strains secreted cellulolytic and xylanolytic enzymes, including endoglucanase, exoglucanase, beta-glucosidase and xylanase. Strain E2 secreted the highest levels of enzymes in a relatively short time. The product formation on avicel by enzymes secreted by the four fungi was studied. Both in the presence and absence of glucurono-1,5-delta-lactone, a specific inhibitor of beta-glucosidase, mainly glucose was formed but no cellobiose. Therefore the exoglucanase secreted by the four fungi is probably a glucohydrolase

Welsch, B.B., 1991. Elephant dentition. Proceedings American Association of Zoo Veterinarians 9.

Wilson, J.F., Mahajan, U., Wainwright, S.A., Croner, L.J., 1991. A continuum model of elephant trunks. J Biomech Eng 113, 79-84.
Abstract: A continuum model is presented that relates the trunk parameters of loading, geometry, and muscle structure to the necessary conditions of static equilibrium. Linear theory for stress-strain behavior is used to describe an elephant trunk for an incremental displacement as the animal slowly lifts a weight at the trunk tip. With this analysis and experimental values for the trunk parameters, the apparent trunk stiffness Ea is estimated for the living animal. For an Asian elephant with a maximum compression strain of 33 percent, Ea is of the order of 10(6) N/m2. The continuum model is quite general and may be applied to similar nonskeletal appendages and bodies of other animals.

Cole, G., Neal, J.W., 1990. The brain in aged elephants. Journal of Neuropathology and Experimental Neurology 49, 190-192.

Fischer, M.S., 1990. The unique ear of elephants and manatees (Mammalia): A phylogenetic paradox. C. R. Acad. Sci. Ser. III Sci. Vie 311, 157-162.

Hattingh, J., Pitts, N.I., Ganhao, M.F., Moyes, D.G., de Vos, V., 1990. Blood constituent responses of animals culled with succinyldicholine and hexamethonium. Journal of the South African Veterinary Medical Association 61, 117-118.
Abstract: Blood constituent responses of elephants and buffaloes culled in the Kruger National Park, using a mixture of succinyldicholine and hexamethonium, were compared to those of animals culled with succinyldicholine only. The results show a decreased physiological response in the animals culled with the mixture, characterized by lower total catecholamine, cortisol and glucose concentrations. Neither a delay of up to 30 min in obtaining blood samples from culled animals, nor a delay of up to 30 min in processing samples obtained immediately after cessation of respiration, gave any significant difference in the blood constituents which were measured.

Rasmussen, L.E.L., Hultgren, B., 1990. Gross and microscopic anatomy of the vomeronasal organ in the Asian elephant. In: McDonald, D.W., Muller-Schwarze, D., Natynczuk, S.E. (Eds.), Chemical signals in vertebrates 5. Oxford University Press, New York, pp. 154-161.

Rasmussen, L.E.L., Munger, B. Micro-anatomy of the trunk tip of Elephas maximus. Chemical Senses 15, 629. 1990.
Ref Type: Abstract
Abstract: Full-text: This study documents the characteristics of the sensory innervation and cutaneous receptors in the dermal and epidermal skin of the extreme trunk tip (finger) and adjacent skin of the Asian elephant Elephas maximus by light microscopy. During the flehmen response the elephant moistens the trunk tip with liquids of interest and apparently uses this tip for transport of such substances to the mucous-filled openings of the incisive ducts, which lead to the vomeronasal organ. We expected to find this region of the trunk tip richly innervated, perhaps with specialized nerve endings, especially in the epidermis. Unexpectedly, our light microscopic examinations demonstrated three distinctive features. First, a uniquely high density of free nerve endings are apparent in the superficial layers of the trunk tip skin. Second, in the skin closely associated with the trunk tip unusual tiny short vibrissal hairs surrounded by hundreds of axons were interspersed with more conventional vibrissal hairs. Third, unique complex branched encapsulated corpuscles were abundant in the superficial layer of the dermis in the area of the tip and in the closely associated skin. This study provides basic histological information about the trunk tip region as the initial part of our investigation of the innervation, cutaneous sensory receptors, especially possible chemosensory receptors of the trunk and its orifices.

Raubenheimer, E.J., Dauth, J., Dryer, M.J., Smith, P.D., Turner, M.L., 1990. Structure and composition of ivory of the African elephant (Loxodonta africanus). South African Journal of Science 86, 192-193.

Roskosz, T., Kobrynczuk, F., 1990. Some reflexions on the shape of foramen magnum in Proboscidea. Annals of Warsaw Agricultural University SGGW AR, Veterinary-Medicine 15, 3-6.

Sreekumar, K.P., Nirmalan, G., 1990. Estimation of the total surface area in Indian elephants (Elephas maximus indicus). Vet. Res. Commun. 14, 5-17.
Abstract: Twenty-four adult Indian elephants (Elephas maximus indicus) of both sexes and different ages and weights, belonging to the Temple Devaswoms, the Forest Department of the Government of Kerala and the Gemini Circus formed the experimental subjects from which formulae were derived to predict the total surface area from either body measurements or areas of individual regions. Several models, using the parameters studied either singly or in combination, were tried independently for males and females and also for adults irrespective of sex. The best prediction of total surface area (S) in m2 was obtained for adults irrespective of sex by using the two parameters, the height at the shoulders (H) in m and forefoot pad circumference (FFC) in m in the formula S = -8.245 + 6.807H + 7.073FFC. No significant improvement in the accuracy of prediction resulted from the use of the independent best fit formulae for males and females. The conventional method of using the exponential of body weight (kg) for predicting surface area was not found to yield an equivalent accuracy in these animals

Stephanos, J.J., Addison, A.W., 1990. Spectroscopic and kinetic aspects of Elephas maximus hemoglobin. Eur. J Biochem. 189, 185-191.
Abstract: In comparison with myoglobin and human and Glycera dibranchiata hemoglobins, the heme distal side amino acid exchanges within the heme environment of elephant tetrameric hemoglobin (Hbe) only slightly affect the electronic and ESR spectra of Hbe(III) and Hbe(II) derivatives, several of which were prepared and characterized by optical and ESR spectroscopy. Addition of 2,3- bisphosphoglycerate or inositol hexakisphosphate to Hbe(II)NO causes tension in the Fe-N(proximal His) bond, although the behaviour differs in detail from that of HbA(II)NO. There are two equilibrium states of Hbe having significantly different kinetics for the Hbe(III)----Hbe(II) reaction of Hbe(III)NO. This autoreduction occurs in the form of two parallel processes, which collapse into one intermediate rate in the presence of Gri(2, 3)P2. The temperature dependences of the rates enable deduction of delta H0 and delta S0 for the linked equilibrium, and yield linear Eyring plots for Hbe(III)NO, from which activation parameters were estimated on the basis of a previously described mechanism

Williams, T.M., 1990. Heat transfer in elephants: thermal partitioning based on skin temperature profiles. Journal of Zoology (Lond) 222, 235-245.
Abstract: The elephant with its low surface-to-volume ratio presents an interesting problem concerning heat dissipation. To understand how such large mammals remain in thermal balance, we determined the major avenues of heat loss for an adult African elephant and an immature Indian elephant. Because conventional physiological measurements are difficult for these animals, the present study used a non-invasive technique, infrared thermography, to measure skin temperatures of each elephant. Detailed surface temperature profiles and surface area measurements of each elephant were used in standard equations for convective, conductive and radiant heat transfer. Results demonstrated that heat transfer by free convection and radiation accounted for 86% of the total heat loss for the elephants at T_a = 12.6 degrees C. Heat transfer across the ears, an important thermal window at high ambient temperatures, represented less than 8% of the total heat loss. Surface area of the animals, and metabolic heat production calculated from total heat loss of the African elephant, scaled predictably with body mass. In contrast, the thermal conductance of the elephants (71.6 W/degree C, African; 84.5 W/degree C, Indian) was three to five times higher than predicted from an allometric relationship for smaller mammals. The high thermal conductance of elephants is attributed to the absence of fur and appears to counteract reduced heat transfer associated with a low surface-to-volume ratio.

Wong, C.L., Wai, M.-K., Cheng, H.-C., Chung, D., Yamashiro, D., 1990. Preliminary study on the antinociceptive effect of elephant beta-endorphin. Clinical and Experimental Pharmacology and Physiology 17, 33-37.
Abstract: 1. Intraventricular administration of human beta-endorphin and elephant beta-endorphin significantly prolonged the tail flick response tested 30 min later. However, elephant beta-endorphin was about 7-8 times more potent than human beta-endorphin in the tail flick test. 2. beta-Endorphin antagonized the antinociceptive effect of both human beta-endorphin and elephant beta-endorphin by the same extent. Naloxone also antagonized the antinociceptive effects of the beta-endorphins but it was less effective than beta-endorphin. 3. Human beta-endorphin and elephant beta-endorphin were of equal potency in inhibiting the abdominal constriction response induced by intraperitoneal (i.p.) acetic acid. Both beta-endorphin and naloxone antagonized these effects of the beta-endorphins with naloxone being more effective. 4. The present study showed that different opioid receptor subtypes may be involved in the tail flick test and the abdominal constriction test. Furthermore, elephant beta-endorphin was a better antinociceptive agent than human beta-endorphin in the tail flick test.

Yu, L.P., La Mar, G.N., Mizukami, H., 1990. Rearrangement of the distal pocket accompanying E7 His----Gln substitution in elephant carbonmonoxy- and oxymyoglobin: ¹H NMR identification of a new aromatic residue in the heme pocket. Biochemistry 29, 2578-2585.
Abstract: Two-dimensional 1H NMR methods have been used to assign side- chain resonances for the residues in the distal heme pocket of elephant carbonmonoxymyoglobin (MbCO) and oxymyoglobin (MbO2). It is shown that, while the other residues in the heme pocket are minimally perturbed, the Phe CD4 residue in elephant MbCO and MbO2 resonates considerably upfield compared to the corresponding residue in sperm whale MbCO. The new NOE connectivities to Val E11 and heme-induced ring current calculations indicate that Phe CD4 has been inserted into the distal heme pocket by reorienting the aromatic side chain and moving the CD corner closer to the heme. The C zeta H proton of the Phe CD4 was found to move toward the iron of the heme by approximately 4 A relative to the position of sperm whale MbCO, requiring minimally a 3-A movement of the CD helical backbone. The significantly altered distal conformation in elephant myoglobin, rather than the single distal E7 substitution, forms a plausible basis for its altered functional properties of lower autoxidation rate, higher redox potential, and increased affinity for CO ligand. These results demonstrate that one-to-one interpretation of amino acid residue substitution (E7 His----Gln) is oversimplified and that conformational changes of substituted proteins which are not readily predicted have to be considered for interpretation of their functional properties

Bianchi, M., 1989. The thickness, shape and arrangement of elastic fibres within the nuchal ligament from various animal species. Anatomischer Anzeiger 169, 53-66.
Abstract: From studies of the nuchal ligament from several species of ruminant (including cattle, sheep, goats, giraffe, dromedary, buffalo), equids (horses, mules, donkeys), carnivores (12 dogs of different breeds and body size) and an elephant, it was concluded that the thickness of the elastic fibres of the ligament was not directly correlated with the size. During postnatal growth the following structural changes were identified: increase in thickness and length of preexisting elastic fibres; progressive increase in the nuber of fibre splittings and of collaterals given off by individual fibres; neoformation of elastin fibres and their addition to existing ones. In bovines, the thickest elastic fibres had attained their full size by 6 months of age.

de Villiers, D.J., Skinner, J.D., Hall-Martin, A.J., 1989. Circulating progesterone concentrations and ovarian functional anatomy in the African elephant (Loxodonta africana). Journal of Reproduction and Fertility 86, 195-201.
Abstract: Mean plasma progesterone concentrations measured in pregnant and non-pregnant elephants did not differ significantly from each other because of considerable variation, particularly for stage of pregnancy. Maximum progesterone values were recorded during early pregnancy (5-8 months) and declined towards term (22 months). The numbers of corpora lutea or total luteal tissue volume were not critical in maintaining progesterone secretion. An increase in plasma progesterone concentrations with the luteal phase of the ovarian cycle was evident. A possible role of the placenta in the second half of gestation is indicated by an increase in fetal progesterone concentrations towards term.

Fujikura, T., Oura, R., Sekine, J., 1989. Comparative morphological studies on digestion physiology of herbivores. 1. Digestibility and particle distribution of digesta and feces of domestic and feral animals. Journal of the Faculty of Agriculture -Tottori-University 25, 87-93.
Abstract: Digesta was collected from a Japanese Black steer 30 months old and feed and faeces samples were collected from a sheep, goat, camel, wallaby, elephant, horse and koala. The composition of feeds and intakes for each animal is given in tables. Digestibilities of particle distribution of DM and acid detergent fibre are discussed and compared between animals.

Harris, C.R. In search of a cervix. Proc.Ann.Elephant Workshop 10. 43-50. 1989.
Ref Type: Conference Proceeding

Langman, V.A., Maloiy, G.M.O., 1989. Passive obligatory heterothermy of the giraffe. J. Physiol. Lond. 415, 89.

Richter, P. Comparative morphology of the thyroid gland of mammals: shape, size, position, blood supply, innervation and histology. Vergleichende, morphologische Studie an der Glandula thyreoidea der Mammalia unter Berucksichtigung von Form, Grosse, Lage, Gefassversorgung, Innervation und histologischem Aufbau. 1-274. 1989. Giessen, Fachbereich Veterinarmedizin, Justus-Liebig-Universitat .
Ref Type: Thesis/Dissertation
Abstract: Literature concerning the thyroid gland in primates, rodents, insectivores, carnivores, artiodactyles, perissodactyles, elephants, lagomorphs, bats and other classes of mammals is reviewed.

Sreekumar, K.P., Nirmalan, G., 1989. Mineral composition of elephant tusks. Indian Journal of Animal Science 59, 1561-1562.

Stone, J., Halasz, P., 1989. Topography of the retina in the elephant Loxodonta africana. Brain,Behavior and Evolution 34, 84-95.
Abstract: The distribution of neurones in the ganglion cell layer of the retina of an African elephant is described. The eye was obtained post-mortem from an infant animal, which died of an unknown disease. It is assumed that most of the neurones observed in the ganglion cell layer are ganglion cells. Ganglion cells concentrate along a horizontal axis extending across the retina inferior to the optic disc, as in the visual streak described in the retina of many mammals. They also concentrate in the upper temporal retina, in a pattern distinctive to elephants. We suggest that this latter concentration has evolved to monitor the animal's trunk. Features of the eye, including its size, orientation and fundal pigmentation, are also described.

Balke, J.M.E., Boever, W.J., Ellersieck, M.R., Seal, U.S., Smith, D.A., 1988. Anatomy of the reproductive tract of the female African elephant (Loxodonta africana) with reference to development of techniques for artificial breeding. Journal of Reproduction and Fertility 84, 485-492.
Abstract: Complete reproductive tracts of 30 female African elephants (5-53 years), obtained during a population reduction procedure, were examined. The reproductive tracts were palpated in situ via the urogenital canal. A plastic speculum (1.3 X 170 cm) was introduced into the canal and dye was injected to simulate the procedure for artificial insemination. The lengths of the reproductive tracts (from the vulva to the ovary) ranged from 120 to 358 cm. The length increased with the size and age of the animal. There was a membranous constriction (hymen) with an orifice, <2cm in diameter, between the urogenital canal and the vagina, in 4 primigravid and in all 13 nulliparous elephants. The vaginal orifice of 13 multiparous elephants consisted of ragged folds of mucous membrane surrounding a single opening, 5-19cm in diameter. The ages at first conception of 4 pregnant elephants with intact hymenal membranes were 10, 12, 13 and 14 years. The hymen was not penetrated as a result of intromission and therefore the site of ejaculation would have been in the urogenital canal of the 4 primigravid elephants.

Balke, J.M.E., Barker, I.K., Hackenberger, M.K., McManamon, R., Boever, W.J., 1988. Reproductive anatomy of three nulliparous Asian elephants: the development of artificial breeding techniques. Zoo Biology 7, 99-113.
Abstract: Detailed gross examinations of the reproductive tracts of three mature female nulliparous Asian elephants were conducted to develop artificial insemination (AI) techniques. Of primary concern was the determination of length characteristics and the size and configuration of the foramina between segments of the tract. The elephants were 13, 28, and 40 years of age and had been maintained in captivity for most of their lives. One elephant died naturally and two were euthanized for health reasons. The reproductive tracts of two of the elephants were manually palpated in situ via the urogenital canal. A fibreoptoscope was used to visualize the internal structures of the terminal reproductive tract of one elephant and to deposit dye into the vagina. The reproductive organs were removed from the body cavity, dissected, measured, and photographed. The major anatomical obstacles to overcome for standard AI procedures (the passage of an AI pipette into the reproductive tract) were the length of the urogenital canal (85-97 cm), the constriction at the urogenital-vaginal junction, and the tight cervix. The reproductive anatomy was compared to that of previous dissections reported in the literature.

Crelin, E.S., 1988. Ligament of the head of the femur in the orangutan and Indian elephant. The Yale Journal of Biology and Medicine 61, 383-389.
Abstract: A literature search revealed that for over 100 years there has been a consensus that the ligament of the head of the femur (LHF) is absent in the orangutan and elephant. A dissection of the hip joints of an adult orangutan and an adult Indian elephant exposed, in each joint, a robust LHF that is functionally important. These LHFs are easily overlooked during a cursory examination of the hip joints because of the way they differ from the human LHF.

Grussen, B. Comparative survey of all literature findings about the anatomy of Indian and African elephants as a basis for practicing veterinary surgeons. Vergleichende Zusammenstellung der Literaturbefunde uber die Anatomie des Indischen und Afrikanischen Elefanten als Grundlage fur tierartzliches Handeln. 1-276. 1988. Hanover.
Ref Type: Report

Henry, R.W., Orosz, S.E., 1988. The muscles of the crus of the African elephant (Loxodonta africana). Anatomia Histologia Embryologia. 17, 370.

Raubenheimer, E.J., Dauth, J., Dreyer, M.J., de Vos, V., 1988. Parotid salivary gland of the African elephant (Loxodonta africana): structure and composition of saliva. Journal of the South African Veterinary Medical Association 59, 184-187.
Abstract: Specimens from parotid salivary glands of full-grown elephant (Loxodonta africana) a (n=6) and saliva aspirated from their main excretory ducts were examined macroscopically and microscopically and analyzed biochemically. The composition of the saliva was compared to that of the blood. The parotids (n=12; mean = 7.4 kg) are homocrine and of a seromucous nature. Myoepithelial cells are well-developed along intercalated ducts and their processes extend to proximal portions of allied acini. The saliva is hypotonic and contains relatively low concentrations of sodium and glucose and high concentrations of potassium, urea, calcium and phosphorus. Absence of detectable levels of alpha-amylase negates a digestive role and the voluminous secrete evidently aids swallowing by moisturizing and lubricating the large mass of ingested leaves, grass and bark.

Easa, P.S., 1987. Chemical composition of the temporal gland secretion of an Asian elephant (Elephas maximus). Elephant 2(3), 67-68.
Abstract: The non-volatile chemical constituents of a temporal gland secretion of a male Asian elephant are reported for the first time, and they seem to be different, in part, from those of African elephant.

Fischer, M.S., 1987. The trunk of elephants. Zeitschrift fur Saugetierkunde 52, 262-263.

Fischer, M.S., Trautmann, U., 1987. Fetuses of African elephants (Loxodonta africana) in photographs. Elephant 2, 40-45.
Abstract: Noticing that almost nothing has been published on the early ontogenetic development in elephants, we want to start to fill this gap by presenting pictures of elephant fetuses. All fetuses are African elephants (Loxodonta africana). Unfortunately, we do not know the age of the fetuses except for that largest one whichis about 8 months old. All specimens were fixated in 4% formalin. The legends will point to the peculiarities in the external morphology of each fetus.

Hackenberger, M.K. Diet digestibilities and ingesta transit times of captive Asian (Elephas maximus) & African (Loxodonta africana) elephants. 1987. Guelph, University of Guelph.
Ref Type: Thesis/Dissertation

Haug, H., 1987. Brain sizes, surfaces, and neuronal sizes of the cortex cerebri: a stereological investigation of man and his variability and a comparison with some mammals (primates, whales, marsupials, insectivores, and one elephant). American Journal of Anatomy 180, 126-142.
Abstract: This study deals with the stereological estimation of macroscopic sizes of brain and cortex, i.e., volume, surface, and folding, and of microscopic neuronal sizes, i.e., density, mean size, size distribution, and number of neurons. The results show that the degree of variability in man amounts to about 15%. A decrease in volume of the different gray structures can be observed in man after the age of 65 years. The surface, folding index, and length of convolution do not alter with aging. The comparison with mammals of various sizes allows the conclusion that there is a high correlation to brain size for nearly all macroscopic values. Man and elephant, however, have a cortical surface which is, in comparison with whales, relatively small. In contrast, whales have very small cortices compared with man. At the cytoarchitectonic level, the neuronal density has a correlation to brain size. Contrary to other mammals, the primates and man have a high fraction of small granular neurons, especially in layer 4. The assumption that the number of cortical neurons beneath a given surface area of cortex is the same in all mammals cannot be verified, especially in those with large brains. The allometric connection between brain size and parameters is not valid for all measurements (e.g., thickness of cortex, mean size of neurons, perikaryal size distribution, and glial density). Yet some other measurements are well correlated.

Lillywhite, H.B., Stein, B.R., 1987. Surface sculpturing and water retention of elephant skin. Journal of Zoology (Lond) 211, 727-734.

Sharma, V.S., Traylor, T.G., Gardiner, R., Mizukami, H., 1987. Reaction of nitric oxide with heme proteins and model compounds of hemoglobin. Biochemistry 26, 3837-3843.
Abstract: Rates for the reaction of nitric oxide with several ferric heme proteins and model compounds have been measured. The NO combination rates are markedly affected by the presence or absence of distal histidine. Elephant myoglobin in which the E7 distal histidine has been replaced by glutamine reacts with NO 500-1000 times faster than do the native hemoglobins or myoglobins. By contrast, there is no difference in the CO combination rate constants of sperm whale and elephant myoglobins. Studies on ferric model compounds for the R and T states of hemoglobin indicate that their NO combination rate constants are similar to those observed for the combination of CO with the corresponding ferro derivatives. The last observation suggests that the presence of an axial water molecule at the ligand binding site of ferric hemoglobin A prevents it from exhibiting significant cooperativity in its reactions with NO.

Hackenberger, M.K., Burton, J.H., Atkinson, J.L., Dickson, K.M. Rate of ingesta passage within captive African elephants (Loxodonta africana). Proc.Ann.Elephant Workshop. 7, 48-58. 1986.
Ref Type: Conference Proceeding

Jongeward, K.A., Marsters, J.C., Mitchell, M.J., Magde, D., Sharma, V.S., 1986. Picosecond geminate recombination of nitrosylmyoglobins. Biochem. Biophys. Res. Commun. 140, 962-966.
Abstract: The kinetics of NO geminate recombination to sperm whale and elephant myoglobins has been studied on the picosecond time scale using an amplified colliding-pulse mode-locked ring dye laser. The dynamics of ligand rebinding are shown to be affected by the distal structure of the protein surrounding the heme pocket.

Mariappa, D., 1986. Anatomy and Histology of the Indian Elephant. Indira Publishing House, Oak Park, MI.
Abstract: This book deals with all aspects of gross anatomy of the Indian elephant. The chapter on histology covers 35 organs. Apart from detailed information on the anatomy and histology of the Indian elephant, the book provides information on many other species of mammals to cater to the needs of veterinarians and comparative anatomists. contemporary thoughts on phylogeny of elephants are also discussed in the introduction.

Meijler, F.L., van der Tweel, L.H., 1986. Electrocardigrams of 10 elephants and a killer whale in Harderwijk. Ned Tijdschr Geneeskd 130, 2344-2348.

Mizukami, H., Bartnicki, D.E., 1986. Unusual myoglobin of elephant. Elephant 2, 80-81.
Abstract: Myoglobins are proteins found in muscle fibers and they store and carry oxygen. They also bind carbon monoxide (CO). Myoglobins of Loxodonta africana and Elephas maximus are different from myoglobins of most other animals. Most significantly, elephant myoglobins react with CO nearly eight times more strongly than other myoglobins. This means that elephants housed close to expressways (where emission of CO from motor vehicles is greatest) would be affected by the toxic gas more than other animals would. On the other hand, elephant myoglobin resists oxidation to a greater extent and, thus, is more stable to the actions of certain toxins.

Mordenti, J., 1986. Man versus beast: pharmacokinetic scaling in mammals. J Pharm Sci 75, 1028-1040.
Abstract: Land mammals range in size from the 3-g shrew to the 3000-kg elephant. Despite this 10(6) range in weight, most land mammals have similar anatomy, physiology, biochemistry, and cellular structure. This similarity has allowed interspecies scaling of physiologic properties such as heart rate, blood flow, blood volume, organ size, and longevity. The equation that is the basis for scaling physiologic properties among mammals is the power equation Y = aWb, where Y is the physiologic variable of interest, W is body weight, and log a is the y-intercept and b is the slope obtained from the plot of log Y versus log W. Animals commonly used in preclinical drug studies (i.e., mice, rats, rabbits, monkeys, and dogs) do not eliminate drugs at the same rate that humans eliminate drugs; small mammals usually eliminate drugs faster than large mammals. Since drug elimination is intimately associated with physiologic properties that are well described among species, it seems reasonable to surmise that drug elimination can be scaled among mammals. Analysis of drug pharmacokinetics in numerous species demonstrates that drug elimination among species is predictable and, in general, obeys the power equation Y = aWb. Early papers on interspecies pharmacokinetic scaling normalized the x- and y-axes to illustrate the superimpossibility of pharmacokinetic curves from different species. More recently, the x- and y-axes have been left in the common units of concentration and time, and individual pharmacokinetic variables have been adjusted to predict pharmacokinetic profiles in an untested species, usually humans.

Rasmussen, L.E., Schmidt, M.J., Daves, G.D., 1986. Chemical communication among Asian elephants. In: Duvall, D., Silverstein, M., Muller-Schwarze, D. (Eds.), Chemical Signals in Vertebrates: Evolutionary, Ecological, and Comparative Aspects. Plenum Press, pp. 627-646.

Yu, N.T., Thompson, H.M., Mizukami, H., Gersonde, K., 1986. The cobalt-nitrosyl stretching vibration as a sensitive resonance Raman probe for distal histidine-nitrosyl interaction in monomeric hemoglobins. Eur. J Biochem. 159, 129-132.
Abstract: The Co-NO stretching vibration has been assigned in the resonance Raman spectra of various cobalt-substituted monomeric hemoglobins by employing isotope-labeling of nitrosyl (14N16O, 15N16O, 14N18O). Monomeric hemoglobins with a distal histidine (sperm whale myoglobin and leghemoglobin) exhibit this vibration at 573-575 cm-1, whereas hemoglobins without distal histidine (elephant myoglobin and insect hemoglobin from Chironomus thummi thummi, CTT III) show this vibration in the range of 553-558 cm- 1. The Fe-NO stretching vibration which occurs in the range of 554-556 cm-1 does not reflect the distal histidine-ligand interaction. Therefore, the Co-NO moiety which is isoelectronic with the Fe-O2 moiety is a good monitor for distal effects on the exogenous ligand of hemoglobins, especially due to the fact that in hemoglobins with distal histidine the Fe-O2 stretching vibration (567-572 cm-1) is similar to the Co-NO stretching vibration

Acharjyo, L.N., Patnaik, S.K., 1985. Appearance of tusks in male Indian elephants (Elephas maximus). Pranikee 6, 86-87.

Alexander, R.M., 1985. The maximum forces exerted by animals. J Exp Biol 115, 231-238.
Abstract: This paper reviews the maximum forces exerted by animals in a wide range of activities including running, jumping, swimming and biting. Most of the data refer to vertebrates and arthropods, ranging in size from 0.5-mg fleas to 3-tonne elephants. Maximum forces exerted on the environment give values of (force/body weight) which lie, in most cases, between 0.5 body mass-1/3 (kg) and 20 body mass-1/3. Maximum forces exerted by major muscle groups give values of (force/body weight) in most cases between 10 body mass-1/3 and 50 body mass-1/3.

Croner, L.J., Wainwright, S.A. Elephant trunks: morphology and motion. American Zoologist 25[4], 12A. 1985.
Ref Type: Abstract
Abstract: Full Text. Soft tissue appendages are biomechanically interesting because they utilize mechanical principles different from those used in appendages with bony frameworks, and because they are versatile. An elephant's trunk is a long tapering structure, nearly circular in cross-section, pierced by two nostrils running up its center, and consisting entirely of soft tissue. Investigation of an embalmed trunk of an Asian elephant confirms that it has four distinct muscle masses -- a radial, a longitudinal, and two oblique layers. Analysis of films taken of an elephant as it lifted a payload with the tip of its trunk indicates that the trunk is capable of shortening at least 30% of its maximum length, and that it has three sections, each of which shortens at a predictable time during a lift. Work in progress analyzes the sequential strain patterns at different parts of a trunk during the performance of other tasks.

Hildebrand, M., Hurley, J.P., 1985. Energy of the oscillating legs of a fast-moving cheetah, pronghorn, jackrabbit, and elephant. J. Morphol. 184, 23-31.
Abstract: Lifelike models of the oscillating legs treated as three-segment systems show the course of kinetic and potential energy over the locomotor cycle for a cheetah, pronghorn, jackrabbit, and elephant running at speeds approaching their maxima. The models can be adjusted to eliminate differences among the animals in time intervals, mass or length of limb, and joint angles. This facilitates analysis of the influence on total energy of each of these variables and of the distribution of mass among leg segments. Fast-cycling legs of the carnivore type have significantly more energy than those of the hoofed type. This may contribute to the lesser endurance that is usual for carnivores that hunt using a high-speed dash

Kerr, E.A., Yu, N.T., Bartnicki, D.E., Mizukami, H., 1985. Resonance raman studies of CO and O₂ binding to elephant myoglobin (distal His(E7)----Gln). Journal of Biological Chemistry 260, 8360-8365.
Abstract: Carbon monoxide and dioxygen were employed as resonance Raman- visible ligands for probing the nature of the heme-binding site in elephant myoglobin, which has glutamine in the distal position (E7) instead of the usual histidine. The distal histidine (E7) residue has been thought to be responsible for weakening carbon monoxide binding to hemoproteins. It is of interest to see how the His(E7)----Gln replacement affects such parameters as nu(Fe-N epsilon), nu(Fe-CO), delta(Fe-C-O), nu(C-O), delta(Fe-O-O), and nu(O-O) vibrational frequencies and relative intensities. Elephant myoglobin has a CO affinity approximately 6 times higher than that for human/sperm whale myoglobin (Mb). If this enhanced affinity were solely due to the removal of some of the steric hindrance that normally tilts the CO off the heme axis, one would expect the nu(Fe-CO) frequency to decrease and the nu(C-O) frequency to increase relative to the corresponding values in sperm whale Mb. However, the opposite was found. In addition, strong enhancement of the Fe-C-O bending mode was observed. These results suggest that the Fe-C-O linkage remains distorted. In elephant Mb, new interactions resulting from the conformational change accompanying ligand binding may be responsible for the increased CO binding. Similar spectra were obtained for elephant and sperm whale oxymyoglobin. This suggests that the interactions of bound O2 are not markedly affected by the glutamine replacement

Meijler, F.L., 1985. Atrioventricular conduction versus heart size from mouse to whale. J Am Coll Cardiol 5 (Pt 2), 363-365.

Scarborough, J., 1985. Galen's dissection of the elephant. Korot 8, 123-134.

Sher, A.V., Garutt, V.E., 1985. New data on the molar morphology of the elephants. Doklady Acad. Nauk SSSR 285, 221-225.

Baranga, J., 1984. The adrenal gland weights of the African elephant, Loxodonta africana. Zeitschrift fur Saugetierkunde 49, 341-348.

Braunitzer, G., Stangl, A., Schrank, B., Krombach, C., Weisner, H., 1984. Phosphate-haemoglobin interaction. The primary structure of the haemoglobin of the African elephant (Loxodonta africana, Proboscidea): asparagine in position 2 of the beta-chain. Hoppe-Seyler's Z. Physiol. Chem. 365, 743-749.
Abstract: The primary structure of the haemoglobin of the African Elephant (Loxodonta africana) is reported. The sequence was determined by means of a sequenator. The haemoglobin differs in 26 amino acids in the alpha-chains in and 27 in the beta-chains from that of adult human hemoglobin. The haemoglobin of the African Elephant, like that of the Indian Elephant and the llama, has only 5 binding sites for polyphosphate. This finding explains the low p(O₂)₅₀ value in whole blood as a result of the lower 2,3-bisphosphoglycerate-haemoglobin interaction. This is discussed in relation to aspects of respiratory physiology; some points are also of interest with regard to the Second Punic War and Hannibal's crossing of the Alps.

Buys, D., Keogh, H.J., 1984. Notes on the microstructure of hair of the Orycteropodidae, Elephantidae, Equidae, Suidae and Giraffidae. South African Journal of Wildlife Research 14, 111-119.
Abstract: The microstructure of hair of seven species of African mammals is described. Distribution notes and micrographs are presented to assist in hair identification.

Hildebrand, M., 1984. Rotations of the leg segments of three fast-running cursors and an elephant. Journal of Mammalogy 65, 718-720.

Krishnamoorthi, R., La Mar, G.N., Mizukami, H., Romero, A., 1984. A proton NMR investigation of the influence of distal glutamine on structural and dynamic properties of elephant metmyoglobin. Journal of Biological Chemistry 259, 265-270.
Abstract: The proton NMR spectra of metmyoglobin from the Asian elephant, which has the replacement of glutamine for the usual distal histidine, are reported and analyzed. In the low pH region, we detect two interconvertible forms of the met-aquo-protein whose relative stabilities are independent of pH, but depend strongly on both temperature and solvent isotope composition. As the pH is raised, both species convert to the met-hydroxy form, as found for other myoglobins. The temperature dependence of the heme methyl shifts for both acidic protein forms indicates essentially high spin character for the iron, and the mean heme methyl shifts are interpreted as indicating one form with a very slightly weaker, and the other with a significantly stronger, axial ligand field than for the unique sperm whale met-aquo-myoglobin. The thermodynamic data for the equilibrium between the two species are consistent with differences of one hydrogen bond between coordinated water and the distal glutamine. Models are proposed where one form of the protein has not only the glutamine carboxyl oxygen acting as a hydrogen-bond acceptor, but also the amine group. We conclude that a distal glutamine can act both as a stronger and as a weaker hydrogen-bond acceptor towards coordinated water than the usual distal histidine. The relative rates of conversion of the two met-aquo-myoglobin forms to MetMbOH is found to be consistent with the proposed structures for the two forms.

Krishnamoorthi, R., La Mar, G.N., Mizukami, H., Romero, A., 1984. A ¹H NMR comparison of the met-cyano complexes of elephant and sperm whale myoglobin. Assignment of labile proton resonances in the heme cavity and determination of the distal glutamine orientation from relaxation data. Journal of Biological Chemistry 259, 8826-8831.
Abstract: The met-cyano complex of elephant myoglobin has been investigated by high field 1H NMR spectroscopy, with special emphasis on the use of exchangeable proton resonances in the heme cavity to obtain structural information on the distal glutamine. Analysis of the distance dependence of relaxation rates and the exchange behavior of the four hyperfine shifted labile proton resonances has led to the assignment of the proximal His-F8 ring and peptide NHs and the His-FG3 ring NH and the distal Gln-E7 amide NH. The similar hyperfine shift patterns for both the apparent heme resonances as well as the labile proton peaks of conserved resonances in elephant and sperm whale met-cyano myoglobins support very similar electronic/molecular structures for their heme cavities. The essentially identical dipolar shifts and dipolar relaxation times for the distal Gln-E7 side chain NH and the distal His-E7 ring NH in sperm whale myoglobin indicate that those labile protons occupy the same geometrical position relative to the iron and heme plane. This geometry is consistent with the distal residue hydrogen bonding to the coordinated ligand. The similar rates and identical mechanisms of exchange with bulk water of the labile protons for the three conserved residues in the elephant and sperm whale heme cavity indicate that the dynamic stability of the proximal side of the heme pocket is unaltered upon the substitution (His----Gln). The much slower exchange rate (by greater than 10(4] of the distal NH in elephant relative to sperm whale myoglobin supports the assignment of the resonance to the intrinsically less labile amide side chain

Krishnamoorthi, R., La Mar, G.N., 1984. Identification of the titrating group in the heme cavity of myoglobin. Evidence for the heme-protein pi-pi interaction. Eur. J Biochem. 138, 135-140.
Abstract: The pH dependence of the proton NMR chemical shifts of met-cyano and deoxy forms of native and reconstituted myoglobins reflects a structural transition in the heme pocket modulated by a single proton with pK 5.1-5.6. Comparison of this pH dependence of sperm whale and elephant myoglobin and that of the former protein reconstituted with esterified hemin eliminates both the distal histidine as well as the heme propionates as the titrating residue. Reconstitution of sperm whale met-cyano myoglobin with hemin modified at the 2,4-positions leads to a systematic variation in the pK for the structural transition, thus indicating the presence of a coupling between the titrating group and the heme pi system. The results are consistent with histidine FG3 (His-FG3) being the titrating group, and a donor-acceptor pi- pi interaction between its imidazole and the heme is proposed.

Poupa, O., Brix, O., 1984. Cardiac beat frequency and oxygen supply: a comparative study. Comp Biochem Physiol A 78, 1-3.
Abstract: The length of diastole in mammals varies between approx 1 s (elephant) and 38 ms (shrew) which makes oxygen supply in high speed cardiac pumps in very small mammals precarious. High capillary density and high blood P50 are reported in mammals with high frequency cardiac cycle. Both are probably insufficient when cardiac frequency is exceedingly high (shrew: 1000 min-1). High respiratory efficiency due to large relative mitochondrial volume per cell (greater than 50%) seems to be preferential solution to maintain sufficient O2-gradient. Similar strategy, i.e. high relative cardiac mitochondrial volume was reported in analogous situation in ice-fish (Chaenocephalus aceratus) where O2 cardiac cell supply is difficult due to the absence of hemoglobin and cardiac myoglobin.

Rowlands, I.W., Weir, B.J., 1984. Mammals: non-primate eutherians. In: Lamming, G.E. (Ed.), Marshall's physiology of reproduction. Churchill Livingstone, New York, pp. 455-658.

Rubin, C.T., Lanyon, L.E., 1984. Dynamic strain similarity in vertebrates; an alternative to allometric limb bone scaling. Journal of Theoretical Biology 107, 321-327.
Abstract: Galileo (1638) observed that "nature cannot grow a tree nor construct an animal beyond a certain size, while retaining the proportions which suffice in the case of a smaller structure". However, subsequent measurement has shown that limb bone dimensions are scaled geometrically with body size (Alexander et al., 1979a), and that the material properties of their constituent bone tissue are similar in animals over a wide range of body weight (Sedlin & Hirsch, 1966; Yamada, 1970; Burstein et al., 1972; Biewener, 1982). If, as suggested in previous scaling arguments (McMahon, 1973; Biewener, 1982), vigorous locomotion involved the same proportional forces over a wide range of animal size, this would create a paradox since large animals would be in far greater danger of skeletal failure than small ones. However, in vivo strain gauge implantations have shown that, during high speed running, axial force as a proportion of body weight (G) in the limb bones of animals decreases as a function of body size from 6.9 G in a 7 kg turkey to 2.8 G in a small (130 kg) horse. Estimates of axial force in larger animals suggest that this is further reduced to 0.8 G in a 2500 kg elephant. Nevertheless, it appears that, regardless of animal size or locomotory style, the peak stresses in the bones of these animals are remarkably similar. Therefore, throughout the range of animals considered (350 times differences in mass), we suggest that similar safety factors to failure are maintained, not by allometrically scaling bone dimensions, but rather by allometrically scaling the magnitude of the peak forces applied to them during vigorous locomotion.(ABSTRACT TRUNCATED AT 250 WORDS)

Wright, P.G., Luck, C.P., 1984. Do elephants need to sweat? South African Journal of Zoology 19, 270-274.

Wright, P.G., 1984. Why do elephants flap their ears? South African Journal of Zoology 19, 266-269.

Bartnicki, D.E., Mizukami, H., Romero-Herrera, A.E., 1983. Interaction of ligands with the distal glutamine in elephant myoglobin. Journal of Biological Chemistry 258, 1599-1602.
Abstract: The effects of distal glutamine (E7) replacement in elephant myoglobin were studied by comparing the temperature-dependent nitrosyl electron spin resonance spectra, redox potentials, and the acid-alkaline equilibria of elephant and human myoglobins. For myoglobins containing a distal histidine, the nitrosyl ESR spectra do not exhibit superhyperfine splitting until near liquid helium temperatures (Yoshimura, T., Ozaki, T., Shintani, Y., and Watanabe, H. (1979) Arch. Biochem. Biophys. 193, 301-313). Studies presented here show that the ESR spectra of nitrosyl elephant myoglobin exhibit 9-line superhyperfine splitting well above liquid nitrogen temperatures, similar to the temperature profiles of isolated heme complexes (Morse, R.H. (1980) Fed. Proc. 39, 2006). It is concluded that the shift in the spectral equilibrium to higher temperature indicates a diminished interaction between NO and the distal position in elephant myoglobin. In addition, the redox potential of elephant myoglobin was found to be nearly 100 mV greater than that of human myoglobin, and the pKa of the acid-alkaline equilibrium (oxidized myoglobin) was 8.5, being 0.4 unit less than that of other vertebrate myoglobins. These different reactivities between elephant and human myoglobins are discussed based on the nature of charge interactions between polar ligands and distal glutamine and histidine

Clemens, E.T., Maloiy, G.M.O., 1983. Nutrient digestibility and gastrointestinal electrolyte flux in the elephant and rhinoceros. Comparative Biochemistry and Physiology [A] 75, 653-658.
Abstract: 1. Nutrient digestibility and absorption-secretion were studied in elephants and rhinoceros. 2. Prehension and diet selection are discussed. 3. Rhinoceros select less fiber, which may account for their greater digestive efficiency. 4. Foregut digestion and fermentation are most evident in the rhinoceros, while elephants possessed greated caecal-colonic digestion. 5. Relative to rhinoceros, elephants demonstrated greater intestinal VFA absorption and less sodium-potassium flux.

Dmytriw, R. Further discussion of an ankle deformity in a young African elephant at the Indianapolis Zoo. AAZPA Annual Conference Proceedings. AAZPA Annual Conference Proceedings , 455-458. 1983.
Ref Type: Conference Proceeding

Heath, E., Jeyendran, R.S., Graham, E.F., 1983. Ultrastructure of spermatozoa of the Asiatic elephant (Elephas maximus). Zbl. Vet. Med. C. Anat. Histol. Embryol. 12, 245-252.
Abstract: A scanning and transmission electron microscopic study of semen collected with the aid of an artificial vagina was carried out. The ultrastructural characteristics of Asiatic bull elephants is compared to that of other mammalian species.

Ruedi, D., Kupfer, U., Girard, J., Gutzwiller, A., 1983. Untersuchungen zur fortflanzungsphysiologie biem Afikanischen elefanten (Loxodonta africana): Samengewihhung bei wildbullen, weitere schritte hinsichtlich kunstlicher besamung. Erkrankungen der Zootiere 341-381.

Valente, A., 1983. Hair structure of the woolly mammoth, Mammuthus primigenius and the modern elephants, Elephas maximus and Loxodonta africanus. Journal of Zoology (Lond) 199, 271-274.
Abstract: The structure of overhairs from a Wooly mammoth, Mammuthus primigenius, 10-13 thousand years old,is compared with that of its living relatives, Elephas maximus and Loxodonta africana. The hair profile, cross-sectional appearance, wholemount and cuticular scale cast of the hairs of the three species were examined and a selected array of photographs representing the hair structure of each species is presented. In general there is little variation between the three species in the gross structure of the overhairs.

Wallach, J.D., Boever, W.J., 1983. Diseases of Exotic Animals. W.B.Saunders, Philadelphis.

Wallach, J.D., Boever, W.J., 1983. Perissodactyla (equids, tapirs, rhinos), Proboscidae (elephants), and Hippopotamidae (hippopotamus). In: Wallach, J.D., Boever, W.J. (Eds.), Diseases of exotic animals. W.B. Saunders Company, Philadelphia, pp. 761-829.

Braunitzer, G., Jelkmann, W., Stangl, A., Schrank, B., Krombach, C., 1982. Hemaglobins, XLVIII: the primary structure of hemoglobin of the Indian elephant (Elephas maximus, Proboscidae): beta 2 = Asn. Hoppe. Seylers. Z. Physiol. Chem. 363, 683-691.
Abstract: The primary structure of the hemoglobin of the Indian Elephant (Elephas maximus) is given. The sequence was determined automatically in a sequenator. By homologous comparison with adult human HbA, the alpha-chains differ by 24 exchanges and the beta-chains by 27 exchanges. Furthermore, we report p(O2)50 values with regard to altered contact sites with 2,3- bisphosphoglycerate in Indian elephant hemoglobin. Our findings explain the low p(O2)50 and the reduced interaction with 2,3- bisphosphoglycerate. Elephant hemoglobin has, like that of the Llama, only five phosphate binding sites. In addition, we have made an attempt to relate these results to aspects of respiratory physiology. Some implications of these biochemical and physiological results, concerning the Second Punic War and Hannibal's Alp transition, are given.

Clemens, E.T., Maloiy, G.M.O., 1982. The digestive physiology of three East African herbivores: the elephant, rhinoceros and hippopotamus. Journal of Zoology (Lond) 198, 141-156.
Abstract: Studies were conducted to compare structural and physiological differences in the digestive functions of three 0. species of large ungulates; the elephant (Loxodonta africana), the Black rhinoceros (Diceros bicornis) and the Hippopotamus (Hippopotamus amphibius). Major differences were noted in the composition of ingesta and the sites of bacterial fermentative activity. Comparisons are also made as to the influence of feeding behaviour on digestive functions, and as to the similarities of their digestive systems to that of domestic animals.

Eltringham, S.K., 1982. Elephants. Blandfort Books,Ltd., United Kingdom.

Foose, T.J. Trophic strategies of ruminant versus nonruminant ungulates. 1982. Chicago, University of Chicago.
Ref Type: Thesis/Dissertation

Hackenberger, M.K., Atkinson, J.L. Digestibility studies with captive Asiatic and African elephants. AAZPA Reg.Conf.Proc. 129-137. 1982.
Ref Type: Conference Proceeding

Harvey, P.H., 1982. On rethinking allometry. Journal of Theoretical Biology 95, 37-41.
Abstract: Analysis of sets of intra-specific and inter-specific allometric relationships reveals than the inter-specific data generally fit an exponential model better than a linear model. The intra-specific data seem equally suited to either model. Skewness of the data and the effect of logarithmic transformations on correlation coefficients are examined in the light of these findings. Inter-species data are approximately lognormally distributed and logarithmic transformations are necessary to produce linear relationships. As a consequence, correlation coefficients usually increase after logarithmic transformation of inter-species data.

Rees, P.S., 1982. Gross assimilation efficiency and food passage time in the African elephant. African Journal of Ecology 20, 193-198.
Abstract: The amount of food consumed and dung voided by two captive African elephants was measured over a period of 7 days. The mean gross assimilation efficiency of the two elephants was calculated to be 22.4%. The food passage time was 21.4 and 46 h for one elephant. Previous estimates of food consumption by wild elephants, based on an earlier, higher estimate of assimilation efficiency, are considered to be too high and have been recalculated.

Shoshani, J., Alder, R., Andrews, K., Baccala, M.J., Barbish, A., Barry, S., Battiata, R., Bedore, M.P., Berbenchuk, S.A., Bielaczyc, R., Booth, G., Bozarth, N., Bulgarelli, M.A., Church, I., Cosgriff, J.W.Jr., Crowe, H., DeFauw, S.L., Denes, L., Efthyvoulidis, E., Ekstrom, M., Engelhard, J.G., English, P., Fairchild, D.Jr., Fisher, C., Frahm, K., Frederick, D., Fried, J., Gaskins, T., Gatt, J., Gentles, W., Goshgarian, H.G., Grabowski, S., Haase, D., Hajj, K., Hall, G., Hawkins, D., Heberer, C., Helinski, A., Henry, S.R., Heyka, C., Hurt, M., Kemppainen, M., Kendra, C., Koenig, J., Konarske, P., Konwinski, S., Kopacz, S., Lakits, V.T., Jr., Lash, S.S., Laughlin, D.C., Meyers, S., Mizeres, N.J., Morehead, K.M., Muraski, A., Murphy, S., Niebala, J., Overbeck, G., Powitz, R., Rafols, J.A., Raymer, S.L., Rezzonica, L., Rossmoore, H.W., Sabo, D., Schwikert, P.J., Shy, E., Skoney, J., Smith, D., Spodarek, K.L., Sujdak, P.J., Tarrant, T., Thielman, R., Tisch, F., Wolowicz, L., Williams, J., Yehiel, D., 1982. On the dissection of a female Asian elephant (Elephas maximus maximus Linnaeus, 1758) and data from other elephants. Elephant 2, 3-93.
Abstract: A 46-year-old female Asian elephant (Elephas maximus maximus Linnaeus, 1758), named "Iki", died on July 8, 1980, at the Ringling Brothers and Barnum & Bailey Circus, Circus World, Haines City, Florida, USA. She was transported to Detroit and was dissected by the Elephant Interest Group (EIG) and friends, Department of Biological Sciences, Wayne State University. The purpose of this continuing study has been to collect data supplemental to that of previous workers, and to enrich knowledge of elephant anatomy, particularly in areas not thoroughly investigated in the past. Some of these findings were compared to those observed in other elephants: "Shirley", "Tulsa", and "Toose" and to the organs of "Ole Diamond" and "Hazel" (see Appendix II).

Jones, R.C., Brosnan, M.F., 1981. Studies of the deferent ducts from the testis of the African elephant, Loxodonta africana. I. Structural differentiation. Journal of Anatomy 132, 371-386.

Jones, R.C., Holt, W.V., 1981. Studies of the deferent ducts from the testis of the African elephant, Loxodonta africana. III. Ultrastructure and cytochemistry of the ductuli efferentes. J Anat 133 (Pt 3), 247-255.
Abstract: The epithelium of the ductuli efferentes is composed of ciliated, principal, halo and basal cells. The supranuclear cytoplasm of ciliated cells is penetrated by particularly long cilial rootlets which are surrounded by numerous elongate mitochondria. Microtubules are arranged along the longitudinal axis of the cells. The spaces between the microvilli of principal cells form canaliculi which penetrate the apical cytoplasm and appear to be involved in endocytotic activity. The supranuclear cytoplasm contains oval mitochondria and numerous vacuoles. Both ciliated and principal cells contain poorly developed Golgi and endoplasmic reticulum, but numerous supranuclear dense bodies are usually present. Supranuclear and basal accumulations of dense bodies were identified as lipofuscin; they were the source of brown pigmentation in the proximal two thirds of the ductuli efferentes. The halo cells were probably macrophages. They occurred quite frequently and contained crescent shaped nuclei and large
accumulations of lipofuscin material.

Paladino, F.V., Spotila, J.R., Pendergast, D. Respiratory variables of Indian and African elephants. American Zoologist 21[4], 1043. 1981.
Ref Type: Abstract
Abstract: Full Text. End expiratory gas samples of Indian and African Elephants were analyzed for O₂ and CO₂. At rest the mean measured O₂ deficit for Adult Indian Elephants was 3.0% O₂ with a CO₂ increment of 3.18% CO₂ (R.Q.=1.06). Immediately after 10 minutes of exercise the 3 adult Indian Elephants had a mean 4.75% O₂ deficit and 5.2% CO₂ increment (R.Q.=1.1). One juvenile Indian Elephant had a resting O₂ deficit of 4.12% and a 4.6% CO₂ increment (R.Q.=1.12) indicating a slightly higher metabolic rate. One adult African Elephant had a resting 4.2% O₂ deficit and a 4.33% CO₂ increment (R.Q.=1.03).

Van Hoven, W., Prins, R.A., Lankhorst, A., 1981. Fermentative digestion in the African elephant. South African Journal of Wildlife Research 11, 78-86.
Abstract: In an investigation into the nature of gastrointestinal fermentation in adult African elephant ( Loxodonta africana L.), the following parameters were determined in various intestinal compartments: volume, pH, concentrations of sugars, bicarbonate, volatile fatty acids (VFA), lactic acid and protozoa. The composition of the digesta was determined by the proximate analysis and the procedure of van Soest. Concentrations of minerals were analyzed in the caecal and colonic fluids. Rates of gas production, composition of the fermentation gas, rates of lactate breakdown, and cellulolytic and amylolytic activity in the gut contents were measured using in vitro incubations. The results show that there is microbial fermentation of sugars, starch and protein from the food in different compartments of the gastrointestinal tract, with most activity occurring in the caecum and colon. Cellulose digestion, however, is surprisingly small. Despite the low rate of fermentation in the gut the elephant is likely to cover a large part of its energy requirement from the metabolism of VFA arising from the hindgut fermentation. In English with Afrikaans summary.

Fowler, M.E., 1980. Hoof, claw, and nail problems in nondomestic animals. Journal of the American Veterinary Medical Association 177, 885-893.

Holt, W.V., Jones, R.C., Skinner, J.D., 1980. Studies of the deferent ducts from the testis of the African elephant, Loxodonta africana. II. Histochemistry of the epididymis. Journal of Anatomy 130, 367-379.
Abstract: The three main segments of the elephant epididymis were examined for the occurrence, in the spermatozoa and lining epithelium, of carbohydrates, neutral lipids and phospholipids, ATPase, alkaline phosphatase, succinic dehydrogenase, glucose-6-phosphate dehydrogenase, diaphorases, hydroxysteroid dehydrogenases, acid phosphatase and non-specific esterase. The most distinct feature of the carbohydrate content of the epididymis was a layer of acidic, alcian blue-positive glycoprotein over the luminal surface of the epithelium, particularly in the terminal segment. PAS-positive, diastase-resistant inclusions were also found throughout the epididymis. Neutral lipid occurred as droplets above and below the nucleus in the epithelium of the middle segment, and as supranuclear accumulations in the terminal segment. All the enzymes except the steroid dehydrogenases were detected in the epididymal epithelium, and all except the steroid dehydrogenases and acid phosphatase were detected in the spermatozoa. There was considerable variation in the intensity of the cytochemical reactions in the epithelium, but not in the spermatozoa, in different regions of the epididymis. In general, the enzymes involved in active transport showed strongest reactions in the initial and terminal segments, the reactions in the stereocilia being the most intense. The enzymes involved in energy metabolism showed strongest reactions in the middle and terminal segments, with the activity being fairly evenly distributed throughout the cytoplasm of the principal cells. However, the two lysosomal enzymes which were studied showed quite different distributions: the reactions for acid phosphatas were strongest in the initial and middle segments, whilst the reactions for non-specific esterase were strongest in the middle and terminal segments. It is suggested that the initial segment is involved in absorptive and anabolic activity, the middle segment in anabolic activity, and the terminal segment (where spermatozoa are stored ready for ejaculation) in considerable metabolic activity and active transport of substrates across the epithelium.

Hooijer, D.A., 1980. Remarks on the dentition and tooth replacement in elephants. Netherlands Journal of Zoology 30, 510-515.
Abstract: It has often been stated that in the modern elephant there is a special kind of forward-moving tooth succession, which would have come in the place of the ordinary vertical tooth replacement in other mammals. This is a mistaken view. In the modern elephant with its disproportionately large cheek teeth vertical tooth replacement has been eliminated because of lack of space, whereas the "horizontal tooth replacement" is more apparent than real. It is basically the same phenomenon that is known in many mammals, Man included, as "closing of the ranks", teeth drifting together so as to assure continuous contact within the functional series. The "horizontal replacement" is not a substitute for the vertical, and is not peculiar to the elephant.

Johnson, D.L., 1980. Problems in the Land Vertebrate Zoogeography of Certain Islands and the Swimming Powers of Elephants. Journal of Biogeography 7, 383-398.
Abstract: The presence of fossil elephants on certain islands off California, in the Mediterranean, in Indonesia, and off China has led to two widely accepted assumptions: (1) that elephants, being poor swimmers, could not have swum to the islands and therefore must have walked to them, which indicates that (2) land bridges once joined the islands to the mainland. These two assumptions have profoundly influenced various insular biogeographic and geologic reconstructions on and around these islands. New evidence, however, shows unequivocally that living elephants are excellent distance swimmers. They swim in a lunging, porpoise-like fashion while using their trunk as a snorkel. Elephant swimming speeds have been measured up to 2.70 km/h, and maximum distances estimated at 48 km. Their chief motives for swimming seem to be that they can see the islands and smell food on them. Because elephants are excellent distance swimmers, we must re-appraise the origin of land vertebrates on all the islands that held elephants, as well as reappraise the geologic reconstructions that assumed land bridges once connected these islands to the mainland.

1979. "Motty" -- Birth of an African/Asian elephant at Chester Zoo. Elephant 1, 36-40.

Alexander, R.M., Jayes, A.S., Maloiy, G.M.O., Wathuta, E.M., 1979. Allometry of the limb bones of mammals from shrews (Sorex) to elephants (Loxodonta). Journal of Zoology (Lond) 189, 305-314.
Abstract: Measurements have been made of the principal leg bones of 37 species representing almost the full range of sizes of terrestrial mammals. The lengths of corresponding bones tend to be proportional to (body mass)^0.35 and the diameters to (body mass)^0.36 except in the family Bovidae in which the exponents for length are much nearer the value of 0.25 predicted by McMahon's (1973) theory of elastic similarity. Comparisons are made between mammals of similar size belonging to different orders.

Alexander, R.M., Maloiy, G.M.O., Hunter, B., Jayes, A.S., Nturibi, J., 1979. Mechanical stresses in fast locomotion of buffalo (Syncerus caffer) and elephant (Loxodonta africana). Journal of Zoology (Lond) 189, 135-144.
Abstract: Films of buffalo and elephant running, and detailed measurements on dissected legs, have been used to estimate the maximum stresses which occur in locomotion, in certain muscles, tendons and bones. These stresses are similar to stresses previously determined for some other, smaller mammals.

Folk, G.E., Folk, M.A., 1979. Physiology of large mammals by implanted radio capsules. In: Amlaner, Jr.C.J., Macdonald, D.W. (Eds.), A handbook on biotelemetry and radio tracking. Pergamon Press, New York, pp. 33-43.

Ommer, P.A., Radhakrishnan, K. Anatomical peculiarties of elephants. State Level Workshop on Elephants. 13-19. 1979. India, College of Veterinary and Animal Sicences, Kerala Agricultural University.
Ref Type: Conference Proceeding

Vendan, C., 1979. The trunk, hand of the elephant. Study of its prehensile and tactile termination. Ann. Chir. Plast. 24, 392-396.

Whitehill, N., 1979. Suggested mechanical model of elephant trunk muscle tissue and its sheer conjecture. Elephant 1, 34-35.

Adams, J., Garcia III, A., Foote, C.S., 1978. Some chemical constituents of the secretion from the temporal gland of the African elephant (Loxodonta africana). Journal of Chemical Ecology 4, 17-25.
Abstract: The temporal glands of African elephants were examined microscopically and histologically, and were found to be modified apocrine sweat glands. The secretion from thse glands was analyzed by gas chromotography and mass spectrometry, and some of the major volatile components have been identified as phenol and m- and p-cresol.

Hass, G., 1978. Behavioural disorders in a female Indian elephant (Elephas maximus bengalensis) with bony structures on the intermediofacial and statoacoustic nerves. Zoologische Garten 48(4,S.), 297-298.

Kozawa, Y., 1978. Comparative histology of proboscidean molar enamel. Kokubyo Gakkai Zasshi 45, 585-606.

De Jong, W.W., Nuy-Terwindt, E.C., Versteeg, M., 1977. Primary structures of alpha crystallin A chains of elephant, whale, hyrax and rhinoceros. Biochim. Biophys. Acta 491, 573-580.
Abstract: As part of a study of the evolutionary development of the eye lens protein alpha-crystallin the 173 residue A chain of this protein has been studied in elephant, whale, hyrax and rhinoceros. The primary sturctures were inferred mainly from amino acid compositions of peptides obtained by enzymatic digestions and CNBr cleavage. The positions of substitutions, as compared to known bovine A chain, were confirmed by Edman degradation. In accordance with the previously observed slow rate of evolution of the A chain only a small number of substitutions were found among these species. Elephant and hyrax share a number of unique substitutions, strongly indicating a common ancestry of these two species within the mammalian class.

Shoshani, J., 1977. General information on elephants with emphasis on tusks. Elephant 1, 20-34.

Bartels, H., 1976. Comparative aspects of respiration and circulation in mammals. Pneumonologie Supplement 1-9.

Dropp, J.J., 1976. Mast cells in mammalian brain. Acta Anat (Basel) 94, 1-21.
Abstract: Mast cells, which had until recently been believed to be not present in the mammalian brain, were studied in the brains of 29 mammalian species. Although there was considerable intraspecific and interspecific variation, mast cells were most numerous within the leptomeninges (especially in those overlying the cerebrum and the dorsal thalamus - most rodents, most carnivores, chimpanzees, squirrel monkeys and elephant), the cerebral cortex (most rodents, tiger, fox, chimpanzee, tarsier, and elephant) and in many nuclei of the dorsal thalamus (most rodents, tiger, lion, and fox). In some mammals, mast cells were also numerous in the stroma of the telencephalic choroid plexuses (chimpanzee, squirrel monkey), the putamen and the claustrum (chimpanzee), the subfornical organ (pack rat, tiger, chimpanzee), the olfactory peduncles (hooded rat, albino rat), the stroma of the diencephalic choroid plexus (lion, chimpanzee, squirrel monkey), the pineal organ (chimpanzee, squirrel monkey), some nuclei of the hypothalamus (tiger), the infundibulum (hooded rat, tiger, fox) the area postrema (pack rat, chinchilla, lion, spider monkey, chimpanzee, fox) and some nuclei and tracts of the metencephalon and the myelencephalon (tiger). Neither the sex of the animal nor electrolytic lesions made in the brains of some of the animals at various times prior to sacrifice appeared to effect the number and the distribution of mast cells. Age-related changes in mast cell number and distribution were detected in the albino rat.

Estes, J.A., Buss, I.O., 1976. Microanatomical structure and development of the African elephants' temporal gland. Mammalia 40, 429-436.
Abstract: The microanatomical structure of temporal glands from thirteen adult African elephants and one fetus, approximately five months old, were examined. The temporal gland consists of compound tubular alveoli interspersed within a loose connective tissue matrix, which is separated into numerous lobes by septa of dense connective tissue. Structurally the gland is nearly identical with the temporal gland of the Asiatic elephant. Metachromatic granules are numerous around the lumenal border of glandular cells suggesting an acid mucopolysaccharide secretory product. Fibroblasts and macrophages are numerous along with occasional plasma cells and mast cells in the loose connective tissue matrix. Mallory's azan indicated that glandular cytoplasm adjacent to the basement membrane was acidophilic whereas the secretion was always basophilic. Frozen sections of temporal glands stained with oil red-O confirmed the presence of lipids throughout the glandular cytoplasm. The presence of myoepithelial cells was suggested, but not confirmed. Histogenesis of the temporal gland at the five month fetal stage indicates epithelial primordia forming downward projections into the connective tissue. Development at this point is similar to that of mammary gland further confirming a common origin with apocrine sweat glands.

Nevill, G.F., Crompton, W.G., Hennessy, M.A., Watson, P.F., 1976. Instrumentation for artificial insemination in the African elephant Loxodonta africana. International Zoo Yearbook 16, 166-171.

1975. Sisson and Grossman's the anatomy of the domestic animals. W.B. Saunders Company, Philadelphia.

Caldwell, J., French, M.R., Idle, J.R., Renwick, A.G., Bassir, O., Williams, R.T., 1975. Conjugation of foreign compounds in the elephant and hyaena. FEBS Lett 60, 391-395.

Cave, A.J.E., 1975. Postcava structure in the elephant and rhinoceros. Journal of Zoology (Lond) 176, 559-565.
Abstract: The mammalian postcava is vulnerable to lumen diminution or collapse under sudden increase of intra-abdominal pressure. Against such collapse its dorsal wall receives an extrinsic protection from the abdominal parietes. Its ventral wall, however, develops an intrinsic protective mechanism in the form of a specialization of its histological architecture. This specialization is most readily noticeable in large-bodied mammals and the details of it are given for four such forms, viz., Asiatic elephant (Elephas), Sumatran rhinoceros (Didermocerus), Black rhinoceros (Diceros) and White rhinoceros (Ceratotherium).

Elder, W.H., Rodgers, D.H., 1975. Body temperature in the African elephant as related to ambient temperature. Mammalia 39, 395-399.

Hiley, P.G., 1975. How the elephant keeps its cool. Natural History 84, 34-41.

Huber, D., Kardum, P., Gomercic, H., 1975. Blood vessels of the fore limb in Indian elephant, Elephas maximus. Veterinarski Arhiv 45, 311-320.

Markowitz, H., Schmidt, M., Nadal, L., Squier, L., 1975. Do elephants ever forget? Journal of Applied Behavior Analysis 8, 333-335.
Abstract: The adult female elephants (Elephas maximus) were tested on a light-dark discrimination problem with an eight year intertrial interval. The first subject took only six minutes to reach criterion and made only 2 errors, suggesting remarkable retention. The other 2 subjects were identified to have visual anomalies which would have gone undetected without this research.

McCullagh, K.G., 1975. Arteriosclerosis in the African elephant: Part 2. Medial sclerosis. Atherosclerosis 21, 37-59.
Abstract: Summary: A type of spontaneous arteriosclerosis, described as medial sclerosis and quite distinct from atherosclerosis, was found in the aortas, coronary arteries and aortic branch arteries of free-living elephants (Loxodonta africana) in Uganda and Kenya. The lesions took the form of calcified fibrotic plaques in the inner tunica media. The calcification appeared to commence in the internal elastic lamina and was associated with atrophy of medial smooth muscle fibres and their replacement by fibrous tissue. In the aorta, medial sclerosis was found to be associated with aortic dilatation, decreased wall thickness and decreased extensibility. These changes were shown to result in substantial increases in the tangential stresses carried by the tissues of the aorta and coronary arteries. As with atherosclerosis, medial sclerosis increased progressively with age; and the approximate involvement of the aorta at different ages could be predicted from linear regression equations. There was no difference in the severity of lesions between male and female animals. Biochemically, the lesions of medial sclerosis were associated with decreased amounts of elastin and increased amounts of collagen in arterial walls. Arterial tissue showing medial calcification always contained less than 30% elastin by weight. In addition, the severity of medial sclerosis in individual elephants was found to be positively correlated with the concentration of calcium in their sera. The pathogenesis of these lesions is discussed and it is suggested that mechanical stress, medial anoxia and high serum calcium levels all contribute to the aetiology of medial sclerosis.

Super, S.J., 1975. Optometric examination of the African elephant, Loxodonta africana africana, in south west Africa. Madogua 9, 45-51.
Abstract: Twenty-one immobilized elephants, optometrically examined in the Etosha National Park, exhibited very little refractive error. "Super Retinoscopy," applying sunlight for external illumination, was used for the first time as a refractive technique. Etorphine hydrochloride (M 99), the immobilizing agent instilled, caused non-reacting miotic pupils. Gross anatomical observations were made on immobilized elephant's eyes as well as on enucleated eyes.

Vogel, J.J., Ennever, J., 1975. Lipid-dependent calcification of elephant tusk matrix. J Dent Res 54, 416.

Walker, E.P., 1975. Order Proboscidea. Mammals of the World. John Hopkins University Press, Baltimore, pp. 1319-1324.

Cmelik, S., Ley, H., 1974. Fatty acid composition of some cellular and subcellular elements of the elephant adrenal gland. Hoppe-Seyler's Z. Physiol. Chem. 355, 797-802.
Abstract: Lipids from the cortex and medulla of the elephant adrenal gland were extracted with chloroform/methanol, and purified over Sephadex G-25. The total lipids, excluding gangliosides were fractionated by a combination of Unisil columns and preparative silica gel G plates and various fractions analyzed by gas-liquid chromatography for their fatty acid components. Results show that nearly 70% of the total lipids in both parts of the gland consist of cholesterol esters differing in the content of polyunsaturated fatty acids. Phospholipids from the cortex and medulla show an identical fatty acid pattern and are characterized by a considerable amount of arachidonic acid. Mitochondrial and microsomal lipids of the whole adrenal gland were extracted, purified and fractionated in the same way. 35% of the mitochondrial lipids consisted of cholesterol esters with C18:1, C16:0 and C20:3(omega 6) as predominant fatty acids. Triglycerides did not contain any higher polyunsaturated fatty acids. In the phospholipids the predominant fatty acids were C18:0, C18:1, C20:4(omega 6) and C16:0. Microsomal lipids yielded 13% cholesterol esters with very few polyunsaturated fatty acids and a considerable amount of C12 and C14 acids. Unlike the mitochondria, microsomes did not contain any triglycerides. Phospholipids of the microsomes were rich in arachidonic acid and contained polyunsaturated acids not present in other fractions. The presence of larger quantities of C16:1 and C16:2 acids in the adrenal gland suggests the possible existence of the omega 7 pathway, although no other metabolites were identified. The fatty acid pattern excludes the possibility of an essential fatty acid deficiency.

Cmelik, S., Ley, H., 1974. Composition of the lipids from the cell sap of the elephant adrenal cortex. Hoppe-Seyler's Z. Physiol. Chem. 355, 1463-1465.

Elder, W.H., Rodgers, D.H., 1974. Immobilization and marking of African elephants and the prediction of body weight from foot circumference. Mammalia 38, 33-53.

Jones, R.C., Skinner, J.D., Rowlands, I.W., 1974. The role of the urogenital ducts of the African elephant, Loxodonta africana. Journal of Reproduction and Fertility 36, 441-442.

Perry, J.S., 1974. Implantation, foetal membranes and early placentation of the African elephant, Loxodonta africana. Philos. Trans. R. Soc. Lond. [Biol] 269, 109-135.

Fawcett, D.W., Neaves, W.B., Flores, M.N., 1973. Comparative observations on intertubular lymphatics and the organization of the interstitial tissue of the mammalian testis. Biology of Reproduction 9, 500-532.

Hanks, J., 1973. Growth and development of the ovary of the African elephant, Loxodonata africana. Puku 7, 126-131.
Abstract: Aspects of growth and development of the ovary of the African elephant are described. There was a pronounced hypertrophy of foetal ovarian interstitial tissue in the second half of gestation. The left ovary was larger than the right in the majority of prepubertal and foetal elephants. There was a gradual increase in the mean number of macroscopically visible follicles from the age of six years up to the mean age of first ovulation at 14 years.

McCullagh, K.G., 1973. Studies on elephant aortic elastic tissue. I. The histochemistry and fine structure of the fiber. Experimental Molecular Pathology 18, 190-201.
Abstract: Elephant arterial elastic laminae were shown to be refractory to staining by orcein or the resorcin dyes, both normally regarded as routine elastic tissue stains. To investigate this more thoroughly, elastin was isolated from the arterial tissue by alkaline hydrolysis and studied in vitro. Compared to elastin from other species, elephant elastin was found to resist alkaline hydrolysis to a greater extent, to possess a greater UV absorption at 275 nm, and to shown an unusual fluorescence at 415 nm. Electron micrographs of elastic fibers in situ demonstrated the presence of large amounts of microbibrillar sheath surrounding the amorphous core. These results are interpreted to indicate the presence, in elephant arterial elastic tissue, of unusually large amounts of a nonelastin component which interferes with the normal staining reactions.

McCullagh, K.G., Derouette, S., Robert, L., 1973. Studies on elephant aortic elastic tissue. II. Amino acid analysis, structural glycoproteins and antigenicity. Experimental Molecular Pathology 18, 202-213.

Siegel, I.M., 1973. Orthotic treatment of tibiotarsal deformity in an elephant. Journal of the American Veterinary Medical Association 163, 544-545.

Dhindsa, D.S., Sedgwick, C.J., Metcalfe, J., 1972. Comparative studies of the respiratory functions of mammalian blood. VIII. Asian elephant (Elephas maximus) and African elephant (Loxodonta africana africana). Respiratory Physiology 14, 332-342.
Abstract: Respiratory characteristics of blood from four Asian and three African elephants were studied. Oxygen dissociation curves of whole blood were constructed at 37 C and corrected to a plasma pH of 7.40. The mean blood P50 values were 25.2 ± 0.5 and 23.2 ± 1.3 mm for Asian and African elephants, respectively, and these values are significantly different (p< 0.01). The Bohr factors for both species were similar and averaged -0.351 ± 0.029 log PO2/ pH. The Haldane effect was similar in both species (5.5 vol% C CO2 at PCO2 = 40 mmHg). The concentration of 2,3-diphosphoglycerate in elephant blood is similar to that found in normal human blood. The blood morphology of both species was similar except that the leukocyte count was significantly higher in Asian elephants. Starch gel electrophoresis showed that hemoglobin of Asian elephants travels at a slower rate than hemoglobin of African elephants, but both migrate faster than human A hemoglobin.

Hanks, J., Short, R.V., 1972. The formation and function of the corpus luteum in the African elephant, Loxodonta africana . Journal of Reproduction and Fertility 29, 79-89.
Abstract: The uterus and ovaries of 617 elephants shot in Zambia were examined. Corpora lutea seem to be necessary for the development of endometrial glands, and before conception can occur, a certain critical mass of luteal tissue has to be achieved by accumulating crops of CL from successive cycles. The elephant can be either monovular or polyovular, and ovulation is spontaneous. New ovulations do not occur during pregnancy, and the presence of an embryo prolongs the life of the CL. There is great variability in luteal size, small CL being commonest in non-pregnant animals and large ones in pregnant animals. The CL do not enlarge during gestation, and some of the smaller ones may regress. The number of CL in pregnant elephants varied with the age of the cow, the younger elephants having a significantly higher number. Larger CL (>20mm in diameter) predominated in older animals. Very little progesterone appears to be secreted by the corpora lutea, and the hormone could not be detected in the peripheral blood during gestation. If progesterone is necessary for pregnancy, the elephant must be extremely sensitive to it, and may be forced to accumulate a large mass of relatively inactive CL before sufficient hormone is available to enable the animal to become pregnant.

Hattingh, J., 1972. A comparative study of transepidermal water loss through the skin of various animals. Comp Biochem Physiol A 43, 715-718.

Haug, H., 1972. The epiphysis and the circumventricular structures of the epithalamus in the brain of the elephant (Loxodonta africana). Zellforsch Mikrosk Anat 129, 533-547.

Maglio, V.J., 1972. Evolution of mastication in the Elephantidae. Evolution 26, 638-658.

McCullagh, K.G., 1972. Arteriosclerosis in the African elephant. I. Intimal artherosclerosis and its possible causes. Atherosclerosis 16, 307-335.

Albl, P., 1971. Studies on assessment of physical condition in African elephants. Biological Conservation 3, 134-140.
Abstract: Series of external measurements were taken from 240 carcasses of African Elephants during the dry season of 1967 in Zambia, in order to investigate fluctuations of subcutaneous fat and muscles. In addition, the ratio of the weight of the kidneys to kidney-fat, and the contents of fat in the bone-marrow, were determined. From these investigations are deduced and described simple criteria for assessment of the physical condition of African Elephants, which criteria allow objective classification of representative population samples. Extensive individual variations of external anatomical features complicate assessment of the condition. Most of the investigated external physical features are more age- than nutrition-dependent. Only the shape of the lumbar region and the kidney-fat index give a fairly reliable indication of the physical condition of the African elephant.

Buss, I.O., Estes, J.A., 1971. The functional significance of movements and position of the pinnae of the African elephant Loxodonta africana. Journal of Mammalogy 52, 21-27.
Abstract: Observations of wild African elephants (Loxodonta africana) in Uganda indicated that flapping and spreading the highly vascularized ears are probably important functions for heat dissipation. Ear flapping increased as ambient temperatures rose and decreased or ceased during cold or rainy weather. Rate of ear flapping was inversely related to wind velocity. Spreading the ears reduced ear flapping, particularly when an elephant faced downwind. Stimuli that elicited alertness, excitement or hostility caused elephants to raise their heads and spread their ears widely and rigidly, and large elephants occasionally flapped their ears loudly and sharply. Flapping and spreading the ears for heat dissipation are generally not interpreted as danger signals by other elephants.

Hanks, J. The reproductive physiology of the African elephant, Loxodonta africana. 1971. University of Cambridge.
Ref Type: Thesis/Dissertation

Lange, W., 1971. Comparative studies on the cerebellum of man, the elephant and certain toothed whales. Verhandlungen der Anatomischen Gesellschaft 65, 137-138.

Montagna, W., 1971. Cutaneous comparative biology. Archives of Dermatology 104, 577-591.
Abstract: This article reports the biological properties and adaptive patterns of structure and function of mammalian skin. There is an inverse relation between the richness of pelage and the thickness and complexity of epidermal undersurface of the epidermis. Active melanocytes are numerous in the viscera and must have other functions than making melanin. The dermis follows a stereotypical structural pattern, but shows many species differences; no animal has the amounts of elastic tissue, vascularity, and nerves as does the human. Hair follicles differ in different species and in various parts of the body, and are parts of the cutaneous sensory system. Sebaceous and apocrine glands secrete pheromones. Except for horses, only man sweats in response to heat stimulation. Thermal sweating, therefore, can be studied only in man.

Pavelka, R., 1971. The peripheral conduction systems in the hindlimb of Elephas maximus. Anatomischer Anzeiger 128, 150-169.

Sikes, S.K., 1971. The Natural History of the African Elephant. American Elsevier Publishing Company, Inc., New York.

Singh, B.S., 1971. Umbilical hernia in an elephant calf. Indian Veterinary Journal 48, 533-536.

Dellenback, R.J., Chien, S., 1970. The extinction coefficient of fibrinogen from man, dog, elephant, sheep and goat at 280 micrometers. Procedings of the Society for Experimental Biology and Medicine 134, 353-355.

Estes, J.A. Observations on the temporal gland of the African elephant. 1970. Washington, USA, Washington State University.
Ref Type: Thesis/Dissertation

Haug, H., 1970. The macroscopic structure of the cerebrum. Qualitative and quantitative studies on the brain of humans, dolphins, and the elephant. Ergebnisse der Anatomie und Entwicklungsgeschichte 43, 3-70.

Haug, H., 1970. Comparative studies of the brains of men, elephants and toothed whales. Verhandlungen der Anatomischen Gesellschaft 64 , 191-195.

Spearman, R.I.C., 1970. The epidermis and its keratinization in the African elephant (Loxodonta africana). Zoologica Africana 5, 327-338.

Elias, H., Haug, H., Lange, W., Schlenska, G., Schwartz, D., 1969. Surface area determination of the cerebral cortex of mammals with special reference to humans, Cetacea, elephants and Marsupialia]. Verh Anat Ges 63, 461-462.

Friant, M., 1969. Brain development and morphology in a proboscidian, the African elephant (Loxodonta africana Blum.). Acta Neurol Psychiatr Belg 69, 20-32.

Haug, H., 1969. On myelinated nerve fibers and myelinated Herring bodies in the neurohypophysis of the elephant. Z Zellforsch Mikrosk Anat 96, 134-141.

Haug, H., 1969. Comparative quantitative studies on brains of man, elephants and some tooth whales. I. Size of the brain cortex. Med Monatsschr 23, 201-205.

Krumrey, W.A., Buss, I.O., 1969. Observations on the adrenal gland of the African elephant. Journal of Mammalogy 50, 90-101.
Abstract: Forty-nine female and 32 male African elephants (Loxodonta africana) were collected from July 19558 to May 1959 in Bunyoro District, Uganda. Reproductive status, body weight, and measurements were recorded and ages estimated. Gross morphology and histology of the adrenal gland are described: histology of the elephant adrenal corresponds with generalized descriptions for those of other eutherian mammals. The fetal cortex is well developed in an elephant 2 months of age, but gradually degenerates and is completely absorbed by the fourth year of postnatal life without giving rise to another transitory zone. No significant increase in relative adrenal weight related to Uganda's December-to-March dry season was detected. Relative adrenal weight of elephants is higher in females than males of comparable age, in immature males that mature males, and in immature females than nonpregnant or nonlactating females. Pronounced increase in relative adrenal weight is related to pregnancy and lactation.

Poczopko, P., 1969. Self-defense against hyperthermia in animals. Acta Physiologica Polonica 20, 893-905.

Duncan, W.R., Garton, G.A., 1968. The fatty acid composition and intramolecular structure of triglycerides from adipose tissue of the hippopotamus and the African elephant. Comp Biochem Physiol 25, 319-325.

Buss, I.O., Johnson, O.W., 1967. Relationships of Leydig cell characteristics and intratesticular testosterone levels to sexual activity in the African elephant. Anatomical Record 157, 191-196.
Abstract: Histological characteristics of testis tissues from 25 African elephants (Loxodonta africana) collected in Uganda, showed no consistent relationships among the following variables: Leydig cell size, cytoplasmic characteristics, and abundance; testicular testosterone content; and age. From these findings, plus field observations of sexual behavior, emerges the hypothesis that individual cyclicity in Leydig cell function was inherent in the elephant population studied. Testosterone content of testes from 32 elephants (including the 25 studied histologically) suggested that lone bulls were not of a senile nature since they contained relatively large quantities of testosterone and were relatively young (from about 12 to 25 years of age). Also, lone bulls were observed searching out estrous females. Among bulls collected from family units and herds, testosterone levels and behavior differed conspicuously. Behavior appeared to be directly related to testosterone content in several instances. Non-aggressive behavior among members of bull herds, plus the high proportion of such individuals with low testosterone content, suggest that some of these animals were in a depressed phase of sexual activity whereas others were undergoing pubertal development.

Johnson, O.W., Buss, I.O., 1967. The testis of the African elephant (Loxodonta africana). II. Development, puberty and weight. Journal of Reproduction and Fertility 13, 23-30.

Johnson, O.W., Buss, I.O., 1967. The testis of the African elephant (Loxodonta africana). I. Histological features. Journal of Reproduction and Fertility 13, 11-21.

Koikegami, H., Ozaki, N., 1967. Studies on the paralimbic brain structures. 3. On the nucleus accumbens and on the olfacto-tegmental tract in the Indian elephant. Acta Med Biol (Niigata) 15, 131-140.

Kubota, K., 1967. Comparative anatomical and neurohistological observations on the tongues of elephants (Elephas indicus and Loxodonta africana). Anatomical Record 157, 505-515.
Abstract: Frozen sections from Indian and African elephant tongues were investigated neurohistologically. On the dorsum there are 3 to 5 vallate papillae. Foliate papillae consisting of 18 to 27 clefts are observed in the posterolateral region of the tongue. Wart-like papillae are distributed along the lateral border of the tongue from the foliate papillae region to the apex. Vallate and foliate papillae contain serous glands but have no taste buds. They are supplied with abundant lamellated corpuscles of Pacinian type in their upper mucosa. The wart-like papillae are composed of two or more papillae, each of which has many secondary papillae supplied with plexiform thin and thick nerves. They bear a few taste buds and contain lamellated corpuscles of Pacinian type. From these neurohistologic characteristics wart-like papillae should be regarded as a receptive organ for secretion of the lingual glands. Lamellated corpuscles of Pacinian type are widely distributed over the whole surface of the tongue. The histologic location of these two structures is of interest in suggesting than they together play important roles as receptors of taste and tactile sensations during mastication of food. Double motor end plates are fround on single muscle fibers. The mixed glands which are plentiful in the inferolateral area of the tongue are in close topographic relation with the wart-like papillae.

Kunzel, E., Luckhaus, G., 1967. Comparative anatomical studies of the soft palate of mammals: the palatal cartilage and the "M. uvulae" of the India elephant (Elephas maximus). Anatomischer Anzeiger 120, 318-322.

Laws, R.M., Parker, I.S.C., Archer, A.L., 1967. Estimating live weights of elephants from hindleg weights. East African Wildlife Journal 5, 106-111.

O'Donoghue, P.N., Sikes, S.K., Turvey, A., 1967. Notes on the adrenal of the African elephant. Journal of Zoology (Lond) 152, 281-286.
Abstract: Sections of the adrenals of two male specimens of Loxodonta africana were examined. Apart from the dimensions of the gland and the amount and disposition of connective tissue in it, the most characteristic features were the very high lipid content of the cortical secretory cells, the large size of these cells, and the occurrence of dark cells in the zona fasciculata.

Rooney, J.R., II, Sack, W.O., Habel, R.F., 1967. Guide to the dissection of the horse. Wolfgang O. Sack, Ithaca, New York.

Short, R.V., Mann, T., Hay, M.F., 1967. Male reproductive organs of the African elephant, Loxodonta africana. Journal of Reproduction and Fertility 13, 517-536.

Young, E., Lombard, C.O., 1967. Physiological values of the African elephant (Loxodonta africana). The Veterinarian 4, 169-172.
Abstract: The following physiological values of African elephants determined in the Kruger National Park are recorded in this article: Red cell count, hematocrit, mean corpuscular volume, mean corpuscular hemoglobin, mean corpuscular hemoglobin concentration, total hemoglobin, erythrocyte sedimentation rate, total white cell count, differential white cell count, blood urea nitrogen, body temperature, and pH of the blood plasma, saliva, urine and faeces.

Haug, H., 1966. Cyto-architectural studies on the cerebral cortex of the elephant. Verh Anat Ges 61, 331-337.

Horstmann, E., 1966. The epidermis of the elephant. Z. Zellforsch. Mikrosk. Anat. 75, 146-59.

Hungerford, D.A., Chandra, H.S., Snyder, R.L., Ulmer, F.A., 1966. Chromosomes of three elephants, two Asian (Elephas maximus) and one African (Loxodonta africana). Cytogenetics 5, 243-246.
Abstract: Cultured somatic cells of three elephants, male and female Asian (Elephas maximus) and female African (Loxodonta africana), had 56 chromosomes. Karyotypes of the Asian specimens were morphologically identical, except for the sex chromosomes, and that of the African specimen closely similar. Identification of the African X was made by inference.

Kamiya, T., Fujita, T., 1966. The intramural pouch in the duodenum of the Indian elephant: a macro- and microscopic study of six cases. Okajimas Folia Anatomica Japonica 42, 281-294.

Laws, R.M., 1966. Age criteria for the African elephant, Loxodonta a. africana. East African Wildlife Journal 4, 1-37.

Buss, I.O., Wallner, A., 1965. Body temperature of the African elephant. Journal of Mammalogy 46, 104-107.

Holm, N.E., 1965. The musculature of the forelimbs of the Elephas indicus. Anatomischer Anzeiger 117, 171-192.

Amoroso, E.C., Perry, J.S., 1964. The foetal membranes and placenta of the African elephant. Philos. Trans. R. Soc. Lond. [Biol] 248, 1-34.

Cave, A.J.E., Aumonier, F.J., 1964. Lymph node structure in an Asiatic elephant. Journal of the Royal Microscopic Society 82, 251-255.

Cooper, R.A., Connell, R.S., Wellings, S.R., 1964. Placenta of the Indian elephant, Elephas indicus. Science 146, 410-412.
Abstract: The placenta of the Indian elephant is incompletely annular and zonary microscopically and occupies the equator of an ovoid chorioallantoic sac. The amnion is fused with the chorion over the zone. Microscopically, the placenta is labyrinthine and endotheliochorial with a rudimentary marginal hematoma. Both macroscopically and microscopically it resembles the placentas of the carnivores, particularly the raccoon, the cat and the dog.

French, J.E., 1964. Atherosclerosis. In: Florey, H. (Ed.), General pathology. Loyd-Luke, London, pp. 418-446.

Jayasinghe, J.B., Fernando, S.D.A., Brito-Babapulle, L.A.P., 1964. The electrocardiogram of a baby elephant. American Heart Journal 67 , 388-390.

Johnson, O.W. Histological and quantitative characteristics of the testes, observations on the teeth and pituitary gland, and the possibility of reproductive cyclicity in the African elephant (Loxodonta africana). 1-124. 1964. Pullman, Washington, USA, Washington State University.
Ref Type: Thesis/Dissertation

Perry, J.S., 1964. The structure and development of the reproductive organs of the female African elephant. Philos. Trans. R. Soc. Lond. [Biol] 248, 35-52.

Villasenor, C., 1964. A new cellular type found in the liver of Elephas maximus. Gaceta Medica de Mexico 94, 403-405.

Bartels, H., Hilpert, P., Barbey, K., Betke, K., Riegel, K., Lang, E.M., Metcalfe, J., 1963. Respiratory functions of blood of the yak, llama, camel, Dybowski deer and African elephant. American Journal of Physiology 205, 331-336.
Abstract: Blood samples from a yak, llama, camel, deer, and African elephant were analyzed for oxygen capacity, "standard bicarbonate" content, oxygen dissociation curve, and the magnitude of the Bohr and Haldane effects. These parameters of the respiratory function of the blood have been related to the morphology of the red cells, to the weights of the animals, and to the important electrolytes in the erythrocytes and in the plasma. The high affinity for oxgen described previously for llama blood is shared by its relative, the camel. Both of these animals have a high concentration of hemoglobin within their erythrocytes. Blood from the African elephant showed the greatest affinity for oxygen among the subjects studied.

Brattstrom, B.H., Stabile, A.J., Williams, F.R., Des Lauiers, J., Pope, D., 1963. Body temperature of Indian elephants. Journal of Mammalogy 44, 282-283.

Dougall, H.W., 1963. On the chemical composition of elephant faeces. East African Wildlife Journal 1, 123.

Engel, S., 1963. The respiratory tissue of the elephant (Elephas indicus), second communication. Acta Anatomica Nipponica 55, 105-111.
Abstract: The acini of the elephant lung are small but extremely numerous, thus providing an extensive respiratory surface. The parenchyma is subdivided by elastic strands encapsulating small areas of parenchyma. Many, especially peripheral, acini have a lymph system of their own, conspicuously marked by round, peduncular lymph nodes. Generally speaking, the Elephant lung is built up according to the usual pattern of the mammalian lung but contains peculiar structures necessary for the bulk of the body and the volume of the lung.

Fernando, S.D.A., Jayasinghe, J.B., Panabokke, R.G., 1963. A study of the temporal gland in an Asiatic elephant (Elephas maximus). Ceylon Veterinary Journal 11, 108-111.

Jayasinghe, J.B., Fernando, S.D.A., Brito-Babapulle, L.A.P., 1963. The electrocardiographic patterns of Elephas maximus -- the elephant of Ceylon. British Veterinary Journal 119, 559-564.

Riggs, A., 1963. The amino acid composition of some mammalian hemoglobins: mouse, guinea pig and elephant. Journal of Biological Chemistry 238, 2983-2987.

Cave, A.J.E., Aumonier, F.J., 1962. Elephant and rhinoceros lymph node histology. Journal of the Royal Microscopic Society 80, 209-214.
Abstract: The histology is described, for the first time, of certain lymph nodes of the African Elephant (Loxodonta africana Blumenbach), of the Great Indian Rhinoceros (Rhinoceros unicornis Linn.) and of the African Black Rhinoceros (Diceros bicornis Linn.): reference is made to the nodes of the African White Rhinoceros (Ceratotherium simum Gray). All nodes studied prove to be haemolymph organs.

Evans, G.H., 1961. Elephants and Their Diseases: A Treatise on Elephants. Government Printing, Rangoon, Burma.

Gill, J., 1960. Rate of passage through the digestive tract of Indian elephants (Elephas maximus) under zoo conditions. Acta Physiologica Polonica 11, 272-289.

Kohira, E., 1960. The spleen of the Elephas indicus. Acta Anatomica Nipponica 35, 253-260.

Shimizu, Y., Fujita, T., Kamiya, T., 1960. Anatomy of a female Indian elephant with special reference to its visceral organs. Acta Anatomica Nipponica 35, 261-301.

Hashimoto, Y., Yamauchi, S., Yasunobo, E., 1956. Dissection of an elephant. Bulletin University Osaka Prefecture series B 6, 30-52.

Shindo, T., Mori, M., 1956. Musculature of the Indian elephant. Part III. Musculature of the trunk, neck, and head. Okajimas Folia Anat. Japonica 29, 17-40.

Shindo, T., Mori, M., 1956. Musculature of the Indian elephant. Part II. Musculature of the hindlimb. Okajimas Folia Anat. Japonica 28, 114-147.

Shindo, T., Mori, M., 1956. Musculature of the Indian elephant. Part I. Musculature of the forelimb. Okajimas Folia Anat. Japonica 28, 89-113.

Friant, M., 1954. [Form of the brain of elephant (Loxodonta africana Blum.) during prenatal life.]
2306. C. R. Hebd. Seances Acad. Sci. 238, 1534-1535.

Hanstrom, B., 1953. The hypophysis in some South African insectivora, carnivora, hyracoidae, proboscidea, artiodactyla and primates. Arkiv foer Zoologi (Stockholm) 4, 187-294.

Hill, W.C.O., 1953. The anatomy of the African elephant. In: Ward, R. (Ed.), The Elephant in East Central Africa. Rowland Ward, Ltd., London and Nairobi, pp. 15-60.

Perry, J.S., 1953. The reproduction of the African Elephant, Loxodonta africana. Philos. Trans. R. Soc. Lond. [Biol] 237, 93-149.

Ayer, A.A., Mariappa, D., 1952. A radiographic study of ossification in the Indian elephant fetus. Journal of the Anatomical Society of India 1, 3-10.

Engel, S., 1952. The respiratory tissue of the elephant (Elephas indicus). Acta Anatomica Nipponica 16 , 308-314.

Friant, M., 1951. [The form of the brain, during ontogeny, in elephants (Elephantidae).]. C. R. Hebd. Seances Acad. Sci. 232, 2137-2138.

Garutt, V.E., 1951. [Modification of the structure of carpal bones of Proboscidea in relation to conditions of environment.]. Dokl. Akad. Nauk SSSR 77, 513-515.

WEGNER, R.N., 1951. [The processes cucularis mandibulae in the Elephantidae, Serenia, Rhinocerotidae and Suidae.]. Anatomischer Anzeiger 98, 66-82.

Ayer, A.A., Mariappa, D., 1950. External characters of three fetuses of the Indian elephant. Proceedings of the Indian Academy of Science XXXI(B), 193-209.

Bourne, G.H., 1949. The mammalian adrenal gland. Oxford at the Clarendon Press, Oxford.

Weatherford, H.L., 1940. Some observations on the tusks of an Indian elephant. The innervation of the pulp. Anatomical Record 76, 81-93.

Wislocki, G.B., 1940. The topography of the hypophysis of the elephant, manatee, and hyrax. Anatomical Record 77, 427-445.

Wislocki, G.B., 1939. Note on the hypophysis of an adult Indian elephant. Anatomical Record 74, 321-328.

Benedict, F.G., Lee, R.C., 1938. Further observations on the physiology of the elephant. Journal of Mammalogy 19, 175-194.

Hill, W.C.O., 1938. The external and radiological anatomy of a foetal Asiatic elephant. Ceylon Journal of Science 21, 31-43.

Hill, W.C.O., 1938. Studies on the cardiac anatomy of the elephant: II -- the heart and great vessels of a foetal Asiatic elephant. Journal of Science 21, 44-61.

King, R.L., Burwell, C.S., White, P.D., 1938. Some notes on the anatomy of the elephant's heart. American Heart Journal 16, 734-743.

Neuville, H., 1937. Recherches comparatives sur l'organe femelle des elephants avec remarques sur les formations dites hymenales. Annales des Scinces Naturelles,Zoologies et Biologie Animale 20 10, 245-295.

Schulte, T.L., 1937. The genito-urinary system of the Elephas indicus male. American Journal of Anatomy 61 , 131-157.

Benedict, F.G., 1936. The physiology of the elephant. Carnegie Institution of Washington, Washington.

Benedict, F.G., Lee, R.C., 1936. Studies on the body temperatures of elephants. Proc Natl Acad Sci U S A 22, 405-408.

Friant, M., 1933. La regression de la levre superieure au cours de l'otogenie individual chez l'elephant. C. R. Acad. Sci. Paris 878-879.

Boyle, D., 1929. Height in elephants. Journal of the Bombay Natural History Society 33, 437.

Eales, N., 1929. The anatomy of a foetal African elephant, Elephas africanas (Loxodonta africana). Part III. The contents of the thorax and abdomen, and the skeleton. Transactions of the Royal Society of Edinburgh 56, part I, 203-246.

Eales, N., 1928. The anatomy of a foetal African elephant, Elephas africanas (Loxodonta africana) Part II. The body muscles. Transactions of the Royal Society of Edinburgh 55, part III, 609-642.

Eales, N., 1926. The anatomy of the head of a foetal African elephant, Elephas africanas (Loxodonta africana). Transactions of the Royal Society of Edinburgh 54, part III, 491-551.

Eales, N.B., 1925. External characteristics, skin, and temporal gland of a foetal African elephant. Procedings of the Zoological Society of London 2, 445-456.

Petit, G., 1924. Sur l'abouchement des canaux deferents et de vesicules seminales au verumontanum de l'elephant. Bull. Mus. Hist. Nat. , Paris 30, 441.

Benedict, F.G., Fox, E.L., Baker, M.L., 1921. The skin temperature of pachyderms. Proceedings of the National Academy of Sciences of the United States of America 7, 154-156.

Putter, A., 1918. Studien uber physiologische Xhnlichkeit. Arch. f. d. ges. Physiol. 172, 367-412.

Bolk, L., 1917. Anatomical notes on a fetus of Elephas africanus. J. Muller, Amsterdam.

Todd, T.W., 1913. Notes on the respiratory system of the elephant. Anatomischer Anzeiger 44, 175-183.

Evans, G.H., 1910. Elephants and Their Diseases: A Treatise on Elephants. Government Printing, Rangoon, Burma.

Assheton, R., Stevens, T.G., 1905. Notes on the structure and the development of the elephant's placenta. Quarterly Journal of Microscopical Science 49, 1-37.

Paterson, A.M., 1898. The genito-urinary organs of the female Indian elephant. Journal of Anatomy and Physiology 12, 582-604.

Beddard, F.E., 1893. On the brain of the African elephant. Procedings of the Zoological Society of London 1893, 311-315.

Smith, F., 1890. The histology of the skin of the elephant. Journal of Anatomy and Physiology 24, 493-503.

Colin, G., 1888. Traite de physiologie comparee des animaux. Paris.
Abstract: Cited in Benedict, 1936.

Anderson, R.J., 1883. A contribution to the anatomy of the Indian elephant. Journal of Anatomy and Physiology 17, 491-494.

Watson, M., 1883. Additional observations on the structure of the female organs of the Indian elephant (Elephas indicus). Procedings of the Zoological Society of London 517-521.

Watson, M., 1881. On the anatomy of the female organs of the Proboscidea. Transactions of the Zoological Society of London 11, 111-130.

Chapman, H.C., 1880. The placenta and generative apparatus of the elephant. Journal of the Academy of Natural Sciences,Philadelphia 8, 413-422.

Forbes, W.A., 1879. On the anatomy of the african elephant (Elephas africanus, Blum.). Procedings of the Zoological Society of London 420-435.

Miall, L.C., Greenwood, F., 1879. The anatomy of the Indian elephant. Part I. The muscles of the extremities. Journal of Anatomy and Physiology 12, 261-287.

Miall, L.C., Greenwood, F., 1879. The anatomy of the Indian elephant. Part II. Muscles of the head and trunk. Journal of Anatomy and Physiology 12, 385-400.

Miall, L.C., Greenwood, F., 1879. The anatomy of the Indian elephant. Part III alimentary canal and its appendages. Journal of Anatomy and Physiology 13, 17-50.

Mojsisovics, A., 1879. Zur kenntris der afrikanischen elefanten. Arch. Naturgesch. 45, 56.

Watson, M., 1875. Contributions to the anatomy of the Indian elephant, Part IV. Muscles and blood vessels of the face and head. Journal of Anatomy and Physiology 9, 118-133.

Watson, M., 1874. Contributions to the anatomy of the Indian elephant (Elephas indicus). Part III. The head. Journal of Anatomy and Physiology 8, 85-94.

Watson, M., 1872. Contributions to the anatomy of the Indian elephant (Elephas indicus), Part II. Urinary and generative organs. Journal of Anatomy and Physiology 7, 60-74.

Watson, M., 1872. Contributions to the anatomy of the Indian elephant. Part I. The thoracic visera. Journal of Anatomy and Physiology 6, 82-94.

Gray, J.E., 1868. Notes on the foetus of an Elephant, etc. Procedings of the Zoological Society of London 491.

Tennent, J.E., 1867. The wild elephant and the method of capturing and taming it in Ceylon. Longmans, Green and Co., London.

Owen, 1857. Description of the foetal membranes and placenta of the elephant (Elephas indicus, Cuv.), with remarks on the value of placentary characteristics in the classification of the Mammalia. Philos. Trans. R. Soc. Lond. [Biol] 147, 347-353.

Gilchrist, W., 1851. A practical treatise on the treatment of the diseases of the elephant, camel and horned cattle, with instructions for preserving their efficiency. Calcutta.
Abstract: Cited in Benedict, 1936.

Harrison, R., 1850. On the larynx, trachea, and oesophagus of the elephant. Proceedings of the Royal Irish Academy 4, 132-135.

Gulliver, G., 1847. Note on the spermatozoa and on the elevator muscles of the penis of the Indian elephant. Procedings of the Zoological Society of London 105.

Mayer, J.F.C., 1847. Beitrage zur anatomie des elefanten und der urbrigen pachydermen. Nova Acta Academia Caesaeae Leopoldino-Carolinae Germanicae Nature Curiosorum 22, 1-88.

J.G.F., 1844. The osteology of the elephant. J. Asiatic Soc. Bengal 13, 915-919.

1839. The Elephant (as he exists in a wild state and as he has been made subservient, in peace and war, to the purposes of man). Harper and Brothers, New York.
Abstract: Note: This work was originally published by the British Society for the Diffusion of Useful Knowledge

Blair, P., 1749. The anatomy of an elephant. The Philosophical Transactions Abridg'd, The Third Edition Corrected 82-169.

Perrault, C., 1734. Description anatomique d'un elephant. Memoires de L'Academie Royal des Sciences 3, 91-156.

Perrault, C., 1734. Memoires pour servir a l'historie naturelle des Animaux. Academie Royal des Sciences, Paris.

Stukeley, W., 1723. Of the spleen, its description and history, to which is added some anatomical observations in the dissection of an elephant. The author, London.

Blair, P., 1710. Osteographia elephantina: or, a full and exact description of all the bones of an elephant which dy'd near Dundee, April the 27th, 1706, with their several dimensions, etc. Part II. Philos. Trans. R. Soc. Lond. [Biol] 27, 117-168.

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