|
.
... |
(The following additional keywords have been used
to categorize articles within this section and may assist your search.)
kidney, renal, urinary,urine, urinalysis
Elephant
Bibliographic
Database
www.elephantcare.org
References updated October 2009 by date of publication, most recent
first.
Brown, J.L.,
Kersey, D.C., Freeman, E.W., Wagener, T., 2009. Assessment of diurnal
urinary cortisol excretion in Asian and African elephants using
different endocrine methods. Zoo. Biol.
Abstract: Longitudinal urine samples were collected from Asian and
African elephants to assess sample processing and immunoassay techniques
for monitoring adrenal activity. Temporal profiles of urinary cortisol
measured by RIA and EIA, with and without dichloromethane extraction,
were similar; all correlation coefficients were >0.90. However, based on
regression analyses, cortisol immunoactivity in extracted samples was
only 72-81% of that of unextracted values. Within assay technique, RIA
values were only 74-81% of EIA values. Collection of 24-hr urine samples
demonstrated a clear diurnal pattern of glucocorticoid excretion, with
the lowest concentrations observed just before midnight and peak
concentrations occurring around 0600-0800 hr. These results indicate
that elephants fit the pattern of a diurnal species, and that
glucocorticoid production is affected by a sleep-wake cycle similar to
that described for other terrestrial mammals. Cortisol can be measured
in both extracted and unextracted urine using RIA and EIA methodologies.
However, unexplained differences in quantitative results suggest there
may be sample matrix effects and that data generated using different
techniques may not be directly comparable or interchangeable. Zoo Biol
28:1-10, 2009. (c) 2009 Wiley-Liss, Inc
Garner, M.M.,
Helmick, K., Ochsenreiter, J., Richman, L.K., Latimer, E., Wise, A.G.,
Maes, R.K., Kiupel, M., Nordhausen, R.W., Zong, J.C., Hayward, G.S.,
2009. Clinico-pathologic features of fatal disease attributed to new
variants of endotheliotropic herpesviruses in two Asian elephants
(Elephas maximus)
119. Vet. Pathol. 46, 97-104.
Abstract: The first herpesviruses described in association with serious
elephant disease were referred to as endotheliotropic herpesviruses (EEHV)
because of their ability to infect capillary endothelial cells and cause
potentially fatal disease. Two related viruses, EEHV1 and EEHV2, have
been described based on genetic composition. This report describes the
similarities and differences in clinicopathologic features of 2 cases of
fatal endotheliotropic herpesvirus infections in Asian elephants caused
by a previously unrecognized virus within the betaherpesvirus subfamily.
EEHV3 is markedly divergent from the 2 previously studied fatal
probosciviruses, based on polymerase chain reaction sequence analysis of
2 segments of the viral genome. In addition to ascites, widespread
visceral edema, petechiae, and capillary damage previously reported,
important findings with EEHV3 infection were the presence of grossly
visible renal medullary hemorrhage, a tropism for larger veins and
arteries in various tissues, relatively high density of renal herpetic
inclusions, and involvement of the retinal vessels. These findings
indicate a less selective organ tropism, and this may confer a higher
degree of virulence for EEHV3
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.
Mason, G.J.,
Veasey, J.S., 2009. How should the psychological well-being of zoo
elephants be objectively investigated?
47. Zoo. Biol.
Abstract: Animal welfare (sometimes termed "well-being") is about
feelings - states such as "suffering" or "contentment" that we can infer
but cannot measure directly. Welfare indices have been developed from
two main sources: studies of suffering humans, and of research animals
deliberately subjected to challenges known to affect emotional state. We
briefly review the resulting indices here, and discuss how well they are
understood for elephants, since objective welfare assessment should play
a central role in evidence-based elephant management. We cover
behavioral and cognitive responses (approach/avoidance; intention,
redirected and displacement activities; vigilance/startle; warning
signals; cognitive biases, apathy and depression-like changes;
stereotypic behavior); physiological responses (sympathetic responses;
corticosteroid output - often assayed non-invasively via urine, feces or
even hair; other aspects of HPA function, e.g. adrenal hypertrophy); and
the potential negative effects of prolonged stress on reproduction (e.g.
reduced gametogenesis; low libido; elevated still-birth rates; poor
maternal care) and health (e.g. poor wound-healing; enhanced disease
rates; shortened lifespans). The best validated, most used welfare
indices for elephants are corticosteroid outputs and stereotypic
behavior. Indices suggested as valid, partially validated, and/or
validated but not yet applied within zoos include: measures of
preference/avoidance; displacement movements; vocal/postural signals of
affective (emotional) state; startle/vigilance; apathy; salivary and
urinary epinephrine; female acyclity; infant mortality rates; skin/foot
infections; cardio-vascular disease; and premature adult death.
Potentially useful indices that have not yet attracted any validation
work in elephants include: operant responding and place preference
tests; intention and vacuum movements; fear/stress pheromone release;
cognitive biases; heart rate, pupil dilation and blood pressure;
corticosteroid assay from hair, especially tail-hairs (to access
endocrine events up to a year ago); adrenal hypertrophy; male
infertility; prolactinemia; and immunological changes. Zoo Biol 28:1-19,
2009. (c) 2009 Wiley-Liss, Inc
Steinetz, B.,
Lasano, S., de Haas van, D.F., Glickman, S., Bergfelt, D., Santymire,
R., Songsassen, N., Swanson, W., 2009. Relaxin concentrations in serum
and urine of endangered and crazy mixed-up species
66. Ann. N. Y. Acad. Sci. 1160, 179-185.
Abstract: The human population explosion has pushed many mammalian
wildlife species to the brink of extinction. Conservationists are
increasingly turning to captive breeding as a means of preserving the
gene pool. We previously reported that serum immunoactive relaxin
provided a reliable means of distinguishing between true and
pseudopregnancy in domestic dogs, and this method has since been found
to be a reliable indicator of true pregnancy in endangered Asian and
African elephants and Sumatran rhinoceroses. Our canine relaxin
radioimmunoassay (RIA) has now been adapted and validated to measure
relaxin in the serum and urine of felids, including domestic and wild
species. Moreover, a commercially available canine serum relaxin kit
(Witness) Relaxin Kit; Synbiotics, San Diego, CA), has been adapted for
reliable detection of relaxin in urine of some felid species. Our
porcine relaxin RIA has also been utilized to investigate the role of
relaxin in reproductive processes of the spotted hyena, a species in
which the female fetuses are severely masculinized in utero. Indeed,
this species might well now be extinct were it not for the timely
secretion of relaxin to enable copulation and birth of young through the
clitoris. Additional studies have suggested relaxin may be a useful
marker of pregnancy in the northern fur seal and the maned wolf (the
former species has been designated as "depleted" and the latter as "near
threatened"). Given appropriate immunoassay reagents, relaxin
determination in body fluids thus provides a powerful tool for
conservationists and biologists investigating reproduction in a wide
variety of endangered and exotic species
Thitaram, C.,
Chansitthiwet, S., Pongsopawijit, P., Brown, J.L., Wongkalasin, W.,
Daram, P., Roongsri, R., Kalmapijit, A., Mahasawangkul, S., Rojanasthien,
S., Colenbrander, B., van der Weijden, G.C., van Eerdenburg, F.J., 2009.
Use of genital inspection and female urine tests to detect oestrus in
captive Asian elephants
116. Anim Reprod. Sci. 115, 267-278.
Abstract: Captive Asian elephant (Elephas maximus) populations are
decreasing due to low birth rates compared to wild elephants. Improving
oestrous detection in female elephants is required to ensure successful
mating in captive and semi-captive herds. Responsive behaviours of eight
semi-captive bull elephants to the uro-genital area (genital inspection
test) or urinary pheromones (urine test) of 14 female elephants
throughout the oestrous cycle were evaluated. Weekly blood samples were
collected for 27 consecutive months (14 months for the genital
inspection test and 13 months for the urine test) from female elephants
to characterize the patterns of circulating progestagen. Responsive
behaviours of bulls were compared between females in the follicular
versus the luteal phase of the cycle. The sensitivity and specificity of
the genital inspection test were 65% and 68%, while those of the urine
test were 52% and 61%, respectively. The bulls showed significantly
higher "genital inspection", "flehmen from genital area" and "trunk on
back" behaviours during the genital inspection test, and "flehmen"
behaviours during the urine test in oestrous than in non-oestrous
females. In sum, this study showed that monitoring sexual behaviours of
Asian elephant bulls towards females or their urine can be used to
detect the oestrous period. Although the sensitivity and specificity of
both tests were not as high as expected, still, these methods appear to
be more efficient at detecting oestrous than traditional methods based
on mahout estimations of female receptivity. The use of genital
inspection and urine tests may lead to more successful matings and thus
to creating self-sustaining populations of captive elephants in range
countries
Hollister-Smith, J.A., Alberts, S.C., Rasmussen, L.E.L., 2008.
Do
male African elephants, Loxodonta africana, signal musth via urine
dribbling?
Animal Behavior 76, 1829-1841.
Abstract:
The
phenomenon of musth in male elephants involves increased sexual
activity, heightened aggression and nearly continuous dribbling of
pungent smelling urine. Urine chemistry during musth is altered,
suggesting that urine may signal the musth status of the individual.
Signalling musth remotely may benefit individuals if it reduces the
likelihood of physical confrontation between males, which can lead to
injury and even death. Few studies, however, have asked whether and how
male elephants respond to urine of other males. We tested two
predictions of the hypothesis that urine signals musth status to male
conspecifics: (1) that male African elephants differentiate musth and
nonmusth urine, and (2) that males differentiate between urine dribbled
during early and late musth. The second prediction stems from the
observation that males lose weight and presumably body condition during
musth. We conducted two related bioassays with 26 captive nonmusth males
ranging from 13 to 52 years of age. In each assay, subjects were
simultaneously presented with three urine samples (nonmusth, early musth,
late musth), each from a different donor male, and a control. We found
that subjects differentiated between musth and nonmusth samples using
their vomeronasal organ system, but did not discriminate between the
samples using their main olfactory system. Males did not differentiate
early from late musth. In addition, we found that subject contextual
factors, specifically age, dominance status and social grouping,
significantly predicted response. We discuss these results within the
framework of male elephant longevity and social relationships and their
importance to reproductive success.
Meyer, J.,
Goodwin, T., Schulte, B., 2008. Intrasexual chemical communication and
social responses ofcaptive female African elephants. Animal Behavior 76,
163-174.
Abstract: In matrilineal societies, competition between females can
occur within and between social units. Dominance hierarchies reduce
costly conflicts when reliable cues of status are available, and
reproductive condition may alter individual or group status. Female
African elephants live in matriarchal groups with linear dominance
hierarchies occurring within and between groups; elephants use chemical
signals to mediate social interactions. If reproductive condition has
important implications for inter- or intragroup behaviour, then females
should discriminate between chemical signal sources that reveal
reproductive condition. We examined whether trunk-tip contacts between
females within a social group were related to phase of oestrus.
Observations were conducted on 21 reproductively viable females at nine
zoological facilities in North America. Females in the follicular phase
received contacts to the urogenital region at a higher rate than did
luteal phase females, and contacts increased with approaching ovulation.
This supports the existence of an oestrous signal. We also examined
whether an oestrous signal was evident by female investigation of urine
collected from the luteal and follicular phases of unfamiliar
conspecifics. Elephants responded to unfamiliar urine more than to the
control, but response rates to the urine types did not differ. Females
within a social unit detected differences in oestrus, but they did not
show such discrimination to urinary signals from unfamiliar females.
Further evaluation of the existence of a female-to-female oestrous
pheromone requires assessing responses to urine from familiar
individuals. Understanding the relationship between oestrous condition
and dominance status can shed light on the adaptive value of sociality.
Rasmssen, H.B.,
Ganswindt, A., Douglas-Hamilton, I., Vollrath, F., 2008. Endocrine and
behavioral changes in male African elephants: Linking hormone changes to
sexual state and reproductive tactics. Hormones and Behavior.
Abstract: Endocrine and behavioral changes in male African elephants:
Linking hormone changes to sexual state and reproductive tactics.Henrik
B Rasmussen, Andre Ganswindt, Iain Douglas-Hamilton, and Fritz
VollrathHormones and Behavior, May 22, 2008
Hormones play a crucial role in mediating genetic and environmental
effects into morphological and behavioral phenotypes. In systems with
alternative reproductive tactics (ART) shifts between tactics are
hypothesized to be under proximate hormonal control. Most studies of the
underlying endocrine changes behind ART have focused on fish and
amphibians rather than mammals and few have investigated the potential
interaction between different endocrine axes in regulating shifts
between conditional dependent tactics. Using a combination of endocrine
and behavioral data from male African elephants we expand on our
previously published analysis and show that the initial increase in
androgens predates the behavioral shifts associated with reproductively
active periods, supporting the role of androgens in activating sexually
active periods in males. A strong interactive effect between androgens
and glucocorticoids was found to determine the presence or absence of
temporal gland secretion and urine dribbling, signals associated with
the competitive reproductive tactic of musth, with elevated
glucocorticoids levels suppressing the occurrence of musth signals. In
addition external environmental conditions affected hormone levels. The
presence of receptive females resulted in elevated androgens in dominant
musth males but increased glucocorticoids in subordinate non-musth
males. The presented data on hormones, behavior and reproductive tactics
strongly support an underlying endocrine mechanism for mediating the
translation of intrinsic as well as extrinsic local conditions into the
conditional dependent reproductive tactics in male elephants via
interactions between the hypothalamic-pituitary-gonadal and -adrenal
axes.
Rasmussen,
H.B., Ganswindt, A., Douglas-Hamilton, I., Vollrath, F., 2008. Endocrine
and behavioral changes in male African elephants: linking hormone
changes to sexual state and reproductive tactics. Horm. Behav. 54,
539-548.
Abstract: Hormones play a crucial role in mediating genetic and
environmental effects into morphological and behavioral phenotypes. In
systems with alternative reproductive tactics (ART) shifts between
tactics are hypothesized to be under proximate hormonal control. Most
studies of the underlying endocrine changes behind ART have focused on
fish and amphibians rather than mammals and few have investigated the
potential interaction between different endocrine axes in regulating
shifts between conditional dependent tactics. Using a combination of
endocrine and behavioral data from male African elephants we expand on
our previously published analysis and show that the initial increase in
androgens predates the behavioral shifts associated with reproductively
active periods, supporting the role of androgens in activating sexually
active periods in males. A strong interactive effect between androgens
and glucocorticoids was found to determine the presence or absence of
temporal gland secretion and urine dribbling, signals associated with
the competitive reproductive tactic of musth, with elevated
glucocorticoids levels suppressing the occurrence of musth signals. In
addition external environmental conditions affected hormone levels. The
presence of receptive females resulted in elevated androgens in dominant
musth males but increased glucocorticoids in subordinate non-musth
males. The presented data on hormones, behavior and reproductive tactics
strongly support an underlying endocrine mechanism for mediating the
translation of intrinsic as well as extrinsic local conditions into the
conditional dependent reproductive tactics in male elephants via
interactions between the hypothalamic-pituitary-gonadal and -adrenal
axes
Slade-Cain,
B.E., Rasmussen, L.E., Schulte, B.A., 2008. Estrous state influences on
investigative, aggressive, and tail flicking behavior in captive female
Asian elephants
78. Zoo. Biol. 27, 167-180.
Abstract: Females of species that live in matrilineal hierarchies may
compete for temporally limited resources, yet maintain social harmony to
facilitate cohesion. The relative degree of aggressive and nonaggressive
interactions may depend on the reproductive condition of sender and
receiver. Individuals can benefit by clearly signaling and detecting
reproductive condition. Asian elephants (Elephas maximus) live in social
matrilineal herds. Females have long estrous cycles (14-16 weeks)
composed of luteal (8-12 weeks) and follicular (4-8 weeks) phases. In
this study, we observed the behavior of four captive Asian elephant
females during multiple estrous cycles over 2 years. We evaluated
whether investigative, aggressive, and tail flicking behaviors were
related to reproductive condition. Investigative trunk tip contacts
showed no distinct pattern by senders, but were more prevalent toward
female elephants that were in their follicular compared with their
luteal phase. The genital area was the most frequently contacted region
and may release reproductively related chemosignals. Aggression did not
differ significantly with estrus; however, rates of aggression were
elevated when senders were approaching ovulation and receivers were in
the luteal phase. Females in the follicular phase may honestly advertise
their condition. Contacts by conspecifics may serve to assess condition
and reduce aggression. A behavior termed "tail flicking" was performed
mainly during the mid-follicular phase when estrogen and luteinizing
hormone levels are known to spike. Tail flicking may disperse chemical
signals in urine or mucus as well as act as a tonic signal that could
provide a means of anticipating forthcoming ovulation by elephants and
also for human observers and caretakers. Zoo Biol 27:167-180, 2008. (c)
2008 Wiley-Liss, Inc
Brown, J.L.,
Somerville, M., Riddle, H.S., Keele, M., Duer, C.K., Freeman, E.W.,
2007. Comparative endocrinology of testicular, adrenal and thyroid
function in captive Asian and African elephant bulls. Gen. Comp
Endocrinol. 151, 153-162.
Abstract: Concentrations of serum testosterone, cortisol, thyroxine
(free and total T4), triiodothyronine (free and total T3) and thyroid
stimulating hormone (TSH) were measured to assess adrenal and thyroid
function as they relate to testicular activity and musth in captive
elephants. Blood samples were collected approximately weekly from Asian
(n=8) and African (n=12) bulls at seven facilities for periods of 4
months to 9.5 years. Age ranges at study onset were 8-50 years for Asian
and 10-21 years for African elephants. Based on keeper logs, seven Asian
and three African bulls exhibited behavioral and/or physical (temporal
gland secretion, TGS, or urine dribbling, UD) signs of musth, which
lasted 2.8+/-2.5 months in duration. Serum testosterone was elevated
during musth, with concentrations often exceeding 100 ng/ml. Patterns of
testosterone secretion and musth varied among bulls with no evidence of
seasonality (P>0.05). Only three bulls at one facility exhibited
classic, well-defined yearly musth cycles. Others exhibited more
irregular cycles, with musth symptoms often occurring more than once a
year. A number of bulls (1 Asian, 9 African) had consistently low
testosterone (<10 ng/ml) and never exhibited significant TGS or UD. At
facilities with multiple bulls (n=3), testosterone concentrations were
highest in the oldest, most dominant male. There were positive
correlations between testosterone and cortisol for six of seven Asian
and all three African males that exhibited musth (range, r=0.23-0.52;
P<0.05), but no significant correlations for bulls that did not
(P>0.05). For the three bulls that exhibited yearly musth cycles, TSH
was positively correlated (range, r=0.22-0.28; P<0.05) and thyroid
hormones (T3, T4) were negatively correlated (range, r=-0.25 to -0.47;
P<0.05) to testosterone secretion. In the remaining bulls, there were no
clear relationships between thyroid activity and musth status. Overall
mean testosterone and cortisol concentrations increased with age for all
bulls combined, whereas thyroid activity declined. In summary, a number
of bulls did not exhibit musth despite being of adequate physical
maturity. Cortisol and testosterone were correlated in most bulls
exhibiting musth, indicating a possible role for the adrenal gland in
modulating or facilitating downstream responses. Data were generally
inconclusive as to a role for thyroid hormones in male reproduction, but
the finding of discrete patterns in bulls showing clear testosterone
cycles suggests they may facilitate expression or control of musth in
some individuals
Brown, J.L.,
Somerville, M., Riddle, H.S., Keele, M., Duer, C.K., Freeman, E.W.,
2007. Comparative endocrinology of testicular, adrenal and thyroid
function in captive Asian and African elephant bulls. General and
Comparative Endocrinology 151, 153-162.
Abstract: Concentrations of serum testosterone, cortisol, thyroxine
(free and total T4), triiodothyronine (free and total T3) and thyroid
stimulating hormone (TSH) were measured to assess adrenal and thyroid
function as they relate to testicular activity and musth in captive
elephants. Blood samples were collected approximately weekly from Asian
(n = 8) and African (n = 12) bulls at seven facilities for periods of 4
months to 9.5 years. Age ranges at study onset were 8-50 years for Asian
and 10-21 years for African elephants. Based on keeper logs, seven Asian
and three African bulls exhibited behavioral and/or physical (temporal
gland secretion, TGS, or urine dribbling, UD) signs of musth, which
lasted 2.8 +/- 2.5 months in duration. Serum testosterone was elevated
during musth, with concentrations often exceeding 100 ng/ml. Patterns of
testosterone secretion and musth varied among bulls with no evidence of
seasonality (P > 0.05). Only three bulls at one facility exhibited
classic, well-defined yearly musth cycles. Others exhibited more
irregular cycles, with musth symptoms often occurring more than once a
year. A number of bulls (I Asian, 9 African) had consistently low
testosterone (< 10 ng/ml) and never exhibited significant TGS or UD. At
facilities with multiple bulls (n = 3), testosterone concentrations were
highest in the oldest, most dominant male. There were positive
correlations between testosterone and cortisol for six of seven Asian
and all three African males that exhibited musth (range, r = 0.23-0.52;
P < 0.05), but no significant correlations for bulls that did not (P >
0.05). For the three bulls that exhibited yearly musth cycles, TSH was
positively correlated (range, r = 0.22-0.28; P < 0.05) and thyroid
hormones (T3, T4) were negatively correlated (range, r = -0.25 to -0.47;
P < 0.05) to testosterone secretion. In the remaining bulls, there were
no clear relationships between thyroid activity and musth status.
Overall mean testosterone and cortisol concentrations increased with age
for all bulls combined, whereas thyroid activity declined. In summary, a
number of bulls did not exhibit musth despite being of adequate physical
maturity. Cortisol and testosterone were correlated in most bulls
exhibiting musth, indicating a possible role for the adrenal gland in
modulating or facilitating downstream responses. Data were generally
inconclusive as to a role for thyroid hormones in male reproduction, but
the finding of discrete patterns in bulls showing clear testosterone
cycles suggests they may facilitate expression or control of musth in
some individuals.
Dehnhard, M.,
2007. Characterisation of the sympathetic nervous system of Asian
(Elephas maximus) and African (Loxodonta africana) elephants based on
urinary catecholamine analyses. Gen. Comp Endocrinol. 151,
274-284.
Abstract: Assessing the welfare status of captive animals using
non-invasive measurements of hormones is of growing interest because
this can serve as an effective tool to facilitate the optimization of
environmental and husbandry conditions. Both the African elephant
(Loxodonta africana) and the Asian elephant (Elephas maximus) exhibit
extremely low breeding success in captivity, and because elevated levels
of stress may negatively influence reproductive functions, this study
sought to establish a method for assessing sympathoadrenal activity in
captive female elephants. We found a circadian variation in urinary
noradrenaline (norepinephrine, NE), adrenaline (epinephrine, Epi) and
dopamine (DA) under short day length. Peak activity of noradrenaline and
dopamine was noted at 3 a.m. Adrenaline showed a biphasic pattern with a
minor peak recorded at 3 a.m. and a major peak 9 a.m. Under long-day
photoperiodic conditions, simultaneous peaks of noradrenaline and
adrenaline were again noted at 3 a.m. whereas dopamine does not appear
to have a distinct circadian pattern under long-day length. A transfer
of two elephant cows resulted in a marked increase in urinary adrenaline
and noradrenaline levels, confirming that the transfer represented a
stressful event. During the peripartal period, noradrenaline
concentrations increased and maximum concentrations were obtained at
delivery. Daily measurements of urinary dopamine throughout the
follicular phase revealed an increase in dopamine secretion close to
ovulation. This increase might indicate a role of dopamine in the
ovulatory mechanisms. These results suggest that changes in urinary
catecholamine excretion reflect fluctuations in sympathoadrenal activity
and may be a useful indicator of stress
Bagley, K.R.,
Goodwin, T.E., Rasmussen, L.E.L., Schulte, B.A., 2006. Male African
elephants, Loxodonta africana, can distinguish oestrous status via
urinary signals. Animal Behaviour 71, 1445.
Abstract: African elephants are a polygynous species that raise
offspring in a matriarchal society. Unlike females, males disperse,
spend time in mate groups and search for mates when mature. Urinary
chemical signals aid males in detecting reproductively active females. A
preovulatory pheromone has been identified in Asian elephants, Elephas
maximus, but has not yet been experimentally identified in African
elephants. In this study, the goal was to determine whether adult
captive male African elephants can distinguish between urine from
conspecific females in luteal and periovulatory oestrous stages as an
indication that a preovulatory pheromone is released in the urine. Urine
was collected from seven different female African elephants during their
luteal and periovulatory periods of oestrus. Bioassays were conducted
with nine adult male elephants housed at six different facilities. Males
were presented with the two urine types and a control sample once a day
over 3 days to reduce sample novelty, which can result in misleadingly
high responses. All mates showed greater chemosensory responses to the
periovulatory urine by trial 3 with the ability to distinguish the
urines increasing over the 3 days. This is the first experimental
behavioural evidence that African elephants release an oestrous
pheromone in the urine. The ability of the captive male elephants to
discern between the two urine types bolsters the hypothesis that there
is a preovulatory pheromone in African elephants and encourages efforts
to identify it.
Goodwin, T.E.,
Eggert, M.S., House, S.J., Weddell, M.E., Schulte, B.A., Rasmussen, L.E.,
2006. Insect pheromones and precursors in female African elephant urine
450. Journal of Chemical Ecology 32, 1849-1853.
Abstract: Using automated solid-phase dynamic extraction and gas
chromatography-mass spectrometry, our search for urinary chemical
signals from ovulatory female African elephants (Loxodonta africana) has
revealed the bark beetle aggregation pheromones frontalin,
exo-brevicomin, and endo-brevicomin, as well as their precursors and the
aphid alarm pheromones (E,E)-alpha-farnesene and (E)-beta-farnesene.
Enantiomeric ratios for brevicomins have been determined. Prior
discovery of common insect/elephant pheromones in Asian elephants,
namely, (Z)-7-dodecenyl acetate and frontalin, suggests that the present
findings may yield valuable insights into chemical communication among
African elephants
Wiedner, E.,
Alleman, R., Isaza, R. Urinalysis in Asian elephants (Elephas maximus).
2006 Proceedings American Association of Zoo Veterinarians. 265. 2006.
Ref Type: Conference Proceeding
Abstract: Elephants have been reported with renal disease, and arthritic
elephants are often maintained on long-term administration of
non-steroidal anti-inflammatory drugs which have been associated with
renal injury in other species. Therefore, establishment of the
reference values of normal urine would be a useful tool to elephant
health care. Routine urinalysis was performed on urine collected from
healthy female adult Asian elephants (Elephas maximus, n=30). Elephants
were included in the study if they had normal serum urea nitrogen and
creatinine serum concentrations, no history of urinary tract disease,
and had not received any medication within the past 3 mo. Urinalysis
included gross description of urine color and clarity, measurement of pH
and specific gravity, biochemical analysis and sediment evaluation.
Presence of protein was assessed by the sulfosalicylic acid tubidimetric
test. All urine samples were submitted for aerobic bacterial culture.
Initial findings included identification of calcium carbonate crystals
in the urine of most of the animals in this study; some (n=5) contained
small numbers of struvite crystals (magnesium ammonium phosphates). Most
samples were alkaline and clarity ranged from clear to flocculent.
Specific gravity tended to be fairly low, and isosthenuria was frequent.
Trace bilirubin was detected in some samples. No ketonuria or
proteinuria was found. Glucosuria, which has been reported as a
transient and normal finding in captive elephants, was not detected in
any samples.
Williams, M.F.,
2006. Morphological evidence of marine adaptations in human kidneys. Med
Hypotheses 66, 247-257.
Abstract: Amongst primates, kidneys normally exhibiting lobulated,
multipyramidal, medullas is a unique attribute of the human species.
Although, kidneys naturally multipyramidal in their medullary morphology
are rare in terrestrial mammals, kidneys with lobulated medullas do
occur in: elephants, bears, rhinoceroses, bison, cattle, pigs, and the
okapi. However, kidneys characterized with multipyramidal medullas are
common in aquatic mammals and are nearly universal in marine mammals. To
avoid the deleterious effects of saline water dehydration, marine
mammals have adaptively thickened the medullas of their kidneys--which
enhances their ability to concentrate excretory salts in the urine.
However, the lobulation of the kidney's medullary region in marine
mammals appears to be an adaptation to expand the surface area between
the medulla and the enveloping outer cortex in order to increase the
volume of marine dietary induced hypertonic plasma that can be
immediately processed for the excretion of excess salts and nitrogenous
waste. A phylogenetic review of freshwater aquatic mammals suggest that
most, if not all, nonmarine aquatic mammals inherited the medullary
pyramids of their kidneys from ancestors who originally inhabited, or
frequented, marine environments. So this suggest that most, if not all,
aquatic mammals exhibiting kidneys with lobulated medullas are either
marine adapted--or are descended from marine antecedents. Additionally,
a phylogenetic review of nonhuman terrestrial mammals possessing kidneys
with multipyramidal medullas suggest that bears, elephants and possibly
rhinoceroses, also, inherited their lobulated medullas from semiaquatic
marine ancestors. The fact that several terrestrial mammalian species of
semiaquatic marine ancestry exhibit kidneys with multipyramidal
medullas, may suggest that humans could have, also, inherited the
lobulated medullas of their kidneys from coastal marine ancestors. And a
specialized marine diet in ancient human ancestry could, also, explain
the reactivation and enumeration of corporeal eccrine sweat glands and
the copious secretion of salt tears. The substantial loss of genetic
variation in humans relative to other hominoid primates, combined with
the apparent isolation of early Pliocene human ancestors from particular
retroviruses that infected all other African primate species, may
suggest that such a semiaquatic marine phase, during the emergence of
Homo, may have occurred on an island off the coast of Africa during the
early Pliocene.
Ganswindt,
A., Rasmussen, H.B., Heistermann, M., Hodges, J.K., 2005. The sexually
active states of free-ranging male African elephants (Loxodonta
africana): defining musth and non-musth using endocrinology, physical
signals, and behavior
652. Horm. Behav. 47, 83-91.
Abstract: Musth in male African elephants, Loxodonta africana, is
associated with increased aggressive behavior, continuous discharge of
urine, copious secretions from the swollen temporal glands, and elevated
androgen levels. During musth, bulls actively seek out and are preferred
by estrous females although sexual activity is not restricted to the
musth condition. The present study combines recently established methods
of fecal hormone analysis with long-term observations on male-female
associations as well as the presence and intensity of physical signals
to provide a more detailed picture about the physical, physiological,
and behavioral characteristics of different states of sexual activity in
free-ranging African elephants. Based on quantitative shifts in
individual bull association patterns, the presence of different physical
signals, and significant differences in androgen levels, a total of
three potential sub-categories for sexually active bulls could be
established. The results demonstrate that elevations in androgen levels
are only observed in sexually active animals showing temporal gland
secretion and/or urine dribbling, but are not related to the age of the
individual. Further, none of the sexually active states showed elevated
glucocorticoid output indicating that musth does not represent an HPA-mediated
stress condition. On the basis of these results, we suggest that the
term "musth" should be exclusively used for the competitive state in
sexually active male elephants and that the presence of urine dribbling
should be the physical signal used for defining this state
Ganswindt,
A., Rasmssen, H.B., Heistermann, M., Hodges, J.K., 2005. The sexually
active states of free-ranging male African elephants (Loxodonta
africana): defining musth and non-musth using endocrinology, physical
signals, and behavior. Horm Behav 47, 83-91.
Abstract: Musth in male African elephants, Loxodonta africana, is
associated with increased aggressive behavior, continuous discharge of
urine, copious secretions from the swollen temporal glands, and elevated
androgen levels. During musth, bulls actively seek out and are preferred
by estrous females although sexual activity is not restricted to the
musth condition. The present study combines recently established methods
of fecal hormone analysis with long-term observations on male-female
associations as well as the presence and intensity of physical signals
to provide a more detailed picture about the physical, physiological,
and behavioral characteristics of different states of sexual activity in
free-ranging African elephants. Based on quantitative shifts in
individual bull association patterns, the presence of different physical
signals, and significant differences in androgen levels, a total of
three potential sub-categories for sexually active bulls could be
established. The results demonstrate that elevations in androgen levels
are only observed in sexually active animals showing temporal gland
secretion and/or urine dribbling, but are not related to the age of the
individual. Further, none of the sexually active states showed elevated
glucocorticoid output indicating that musth does not represent an HPA-mediated
stress condition. On the basis of these results, we suggest that the
term "musth" should be exclusively used for the competitive state in
sexually active male elephants and that the presence of urine dribbling
should be the physical signal used for defining this state.
Ganswindt,
A., Heistermann, M., Hodges, K., 2005. Physical, physiological, and
behavioral correlates of musth in captive African elephants (Loxodonta
africana)
594. Physiol Biochem. Zool. 78, 505-514.
Abstract: Although musth in male African elephants (Loxodonta africana)
is known to be associated with increased aggressiveness, urine dribbling
(UD), temporal gland secretion (TGS), and elevated androgens, the
temporal relationship between these changes has not been examined. Here,
we describe the pattern of musth-related characteristics in 14 captive
elephant bulls by combining long-term observations of physical and
behavioral changes with physiological data on testicular and adrenal
function. The length of musth periods was highly variable but according
to our data set not related to age. Our data also confirm that musth is
associated with elevated androgens and, in this respect, show that TGS
and UD are downstream effects of this elevation, with TGS responding
earlier and to lower androgen levels than UD. Because the majority of
musth periods were associated with a decrease in glucocorticoid levels,
our data also indicate that musth does not represent a physiological
stress mediated by the hypothalamic-pituitary-adrenal axis. Furthermore,
we demonstrate that the occurrence of musth is associated with increased
aggression and that this is presumably androgen mediated because
aggressive males had higher androgen levels. Collectively, the
information generated contributes to a better understanding of what
characterizes and initiates musth in captive African elephants and
provides a basis for further studies designed to examine in more detail
the factors regulating the intensity and duration of musth
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
Rasmussen,
L.E.L., Krishamurthy, V., Sakumar, R., 2005. Behavioural and chemical
confirmation of the preovulatory pheromone, (Z)-7-dodecenyl acetate, in
wild Asian elephants: its relationship to musth. Behaviour 142,
351-396.
Abstract: Mammalian breeding strategies vary depending on particular
social contexts and sensory systems emphasized in various species. Among
sexually dimorphic non-territorial Asian elephants, Elephas maximus,
a multiplex olfactory chemical signaling system has been implicated in
ensuring effective reproduction. This study explores how, using
chemosensory mechanisms, widely roaming, wild male elephants locate
periovulatory females in matriarchal-led female family units and
precisely assess their ovulatory status. In this species, the dual
obstacles of separately living sexes and infrequent oestrus are overcome
by lengthy female cycles. During an extended preovulatory period captive
females release increasing concentrations of the urinary pheromone
(Z)-7-dodecenyl acetate, timed to reach a maximum just before ovulation.
The current field studies combined chemical identification and
quantification of female urinary (Z)-7-dodecenyl acetate with
behavioural observations, monitoring the frequencies of chemosensory
responses and premating behaviours by various categories of males. The
results suggest the temporal extension of the preovulatory period
effectively provides a synchrony between sexes for successful
reproduction. Male elephants undergo a two-decade-long maturation
process that involves physical, sexual, social, and physiological
maturation. Males older than 30 years are generally large, sexually
active, socially adept and capable of sustaining long periods of musth,
during which they release secretions distinctive of adult musth. These
older adult males in musth demonstrated significantly more chemosensory
responses and premating behaviours than their younger or nonmusth
counterparts; they apparently are more skilled at detecting the precise
ovulatory status of females. Male-male interactions are affected by
size, age, and musth; the winners gain greater access to females, as
indicated by the high incidence of mate guarding. The Asian elephant
shares some breeding tactics common to other mammals including some
primates (e.g. orangutans) and whales, while the musth parameter adds a
unique feature. Fusion-fission events are influenced by elephant
reproductive strategies, as roving males join female groups while
tracking preovulatory pheromone concentrations.
Steinetz, B.G.,
Brown, J.L., Roth, T.L., Czekala, N., 2005. Relaxin concentrations in
serum and urine of endangered species: correlations with physiologic
events and use as a marker of pregnancy
596. Ann. N. Y. Acad. Sci. 1041, 367-378.
Abstract: Many mammalian species are facing extinction due to problems
created by human encroachment, agriculture, pollution, and willful
slaughter. Among those at risk are the Asian and African elephant,
Sumatran rhinoceros, and giant panda. Conservation groups try to save
species in the wild by preserving habitat and limiting animal-human
conflicts, often with limited success. Another alternative is to
preserve the extant gene pool through captive breeding as a hedge
against extinction. Measurement of circulating reproductive hormones is
impractical for most wildlife species; determination of urinary or fecal
hormone metabolites provides a more viable approach. To aid breeding
management, one important tool is the ability to diagnose and monitor
pregnancy, especially in species with long gestations (e.g., rhinos over
15 mo and elephants over 20 mo). Unfortunately, measuring progestins
often is not useful diagnostically, because concentrations are similar
during at least part of the pregnancy and the nonpregnant luteal phase
in some species (e.g., elephants, rhinoceroses, and giant pandas). As
serum relaxin reliably distinguishes between pregnancy and
pseudopregnancy in bitches, relaxin measurement might also provide a
method for detecting a successful pregnancy in endangered species.
Appropriate immunoassay reagents have enabled the estimation of relaxin
concentrations in the serum of elephants and rhinos and the
determination of pregnancy establishment and the outcome. Relaxin was
also detected in panda serum and urine. However, the extreme variability
of the time between observed mating and parturition and the confounding
factors of delayed implantation, pseudopregnancy, and frequent fetal
resorptions made it impossible to use the panda relaxin data as a
specific marker of pregnancy
Brown, J.L.,
Olson, D., Keele, M., Freeman, E.W., 2004. Survey of the reproductive
cyclicity status of Asian and African elephants in North America. Zoo
Biology 23, 309-321.
Abstract: The Asian and African elephant populations in North America
are not self sustaining, and reproductive rates remain low. One problem
identified from routine progestagen analyses is that some elephant
females do not exhibit normal ovarian cycles. To better understand the
extent of this problem, the Elephant TAG/SSP conducted a survey to
determine the reproductive status of the captive population based on
hormone and ultrasound evaluations. The survey response rates for
facilities with Asian and African elephants were 81% and 71%,
respectively, for the studbook populations, and nearly 100% for the SSP
facilities. Of the elephants surveyed, 49% of Asian and 62% of African
elephant females were being monitored for ovarian cyclicity via serum or
urinary progestagen analyses on a weekly basis. Of these, 14% of Asian
and 29% of African elephants either were not cycling at all or exhibited
irregular cycles. For both species, ovarian inactivity was more
prevalent in the older age categories (>30 years); however, acyclicity
was found in all age groups of African elephants. Fewer elephant females
(B30%) had been examined by transrectal ultrasound to assess
reproductive-tract integrity, and corresponding hormonal data were
available for about three-quarters of these females. Within this subset,
most (B75%) cycling females had normal reproductive-tract morphologies,
whereas at least 70% of noncycling females exhibited some type of
ovarian or uterine pathology. In summary, the survey results suggest
that ovarian inactivity is a significant reproductive problem for
elephants held in zoos, especially African elephants. To increase the
fecundity of captive elephants, females should be bred at a young age,
before reproductive pathologies occur. However, a significant number of
older Asian elephants are still not being reproductively monitored. More
significantly, many prime reproductive-age (10-30 years) African females
are not being monitored. This lack of information makes it difficult to
determine what factors affect the reproductive health of elephants, and
to develop mitigating treatments to reinitiate reproductive cyclicity.
Brown, J.L.,
Goritz, F., Pratt-Hawkes, N., Hermes, R., Galloway, M., Graham, L.H.,
Gray, C., Walker, S.L., Gomez, A., Moreland, R., Murray, S., Schmitt,
D.L., Howard, J., Lehnhardt, J., Beck, B., Bellem, A., Montali, R.,
Hildebrandt, T.B., 2004. Successful artificial insemination of an Asian
elephant at the National Zoological Park. Zoo Biology 23, 45-63.
Abstract: For decades, attempts to breed elephants using artificial
insemination (AI) have failed despite considerable efforts and the use
of various approaches. However, recent advances in equipment technology
and endocrine-monitoring techniques have resulted in 12 elephants
conceiving by AI within a 4-year period (19982002). The successful AT
technique employs a unique endoscope-guided catheter and transrectal
ultrasound to deliver semen into the anterior vagina or cervix, and uses
the "double LH surge" (i.e., identifying the anovulatory LH (anLH) surge
that predictably occurs 3 weeks before the ovulatory LH (ovLH) surge to
time insemination. This study describes the 6-year collaboration between
the National Zoological Park (NZP) and the Institute for Zoo Biology and
Wildlife Research (IZW), Berlin, Germany, that led to the refinement of
this AI technique and subsequent production of an Asian elephant calf.
The NZP female was the first elephant to be inseminated using the new AI
approach, and was the fifth to conceive. A total of six AI trials were
conducted beginning in 1995, and conception occurred in 2000. Semen was
collected by manual rectal stimulation from several bulls in North
America. Sperm quality among the bulls was variable and was thus a
limiting factor for AI. For the successful AI, semen quality was good to
excellent (75-90% motile sperm), and sperm was deposited into the
anterior vagina on the day before and the day of the ovLH surge. Based
on transrectal ultrasound, ovulation occurred the day after the ovLH
surge. Pregnancy was monitored by serum and urinary progestagen, and
serum prolactin analyses in samples collected weekly. Fetal development
was assessed at 12, 20, and 28 weeks of gestation using transrectal
ultrasound. Elevated testosterone measured in the maternal circulation
after 36 weeks of gestation reliably predicted the calf was a male.
Parturition was induced by administration of 40 IU oxytocin 3 days after
serum progestagens dropped to undetectable baseline levels. We conclude
that AI has potential as a supplement to natural breeding, and will be
invaluable for improving the genetic management of elephants, provided
that problems associated with inadequate numbers of trained personnel
and semen donors are resolved.
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
Sanchez,
C.R., Murray, S., Montali, R.J., Spelman, L.H., 2004. Diagnosis and
treatment of presumptive pyelonephritis in an Asian elephant (Elephas
maximus). J Zoo Wildl Med 35, 397-399.
Abstract: A 37-yr-old female Asian elephant (Elephas maximus) presented
with anorexia, restlessness, and dark-colored urine. Urinalyses showed
hematuria, leukocyturia, isosthenuria, proteinuria, granular casts, and
no calcium oxalate crystals. Bloodwork revealed azotemia. Urine culture
revealed a pure growth of Streptococcus zooepidemicus resistant to
sulfamethoxazole-trimethoprim but susceptible to cephalosporins. A
presumptive diagnosis of pyelonephritis was made based on bloodwork,
urinalysis, and urine culture. The animal was treated with intravenous
ceftiofur, and intravenous and per rectum fluids were given for
hydration. The elephant's attitude and appetite returned to normal, the
abnormal blood parameters resolved, and urinary calcium oxalate crystals
reappeared after treatment, supporting presumptive diagnosis. Follow-up
ultrasonography revealed an abnormal outline of both kidneys with
parenchymal hyperechogenicity and multiple uterine leiomyomas.
Suarez, R.K.,
Darveau, C.A., Childress, J.J., 2004. Metabolic scaling: a many-splendoured
thing
656. Comp Biochem. Physiol B Biochem. Mol. Biol. 139, 531-541.
Abstract: Animals at rest and during exercise display rates of aerobic
metabolism, VO2, that represent mainly the sum of mitochondrial
respiration rates in various organs. The relative contributions of these
organs change with physiological state such that internal organs such as
liver, kidney and brain account for most of the whole-body VO2 at rest,
while locomotory muscles account for >90% of the maximum rate, VO2max,
during maximal aerobic exercise. Mechanisms that regulate VO2 are
complex and the relative importance of each step in a series, estimated
by metabolic control analysis, depends upon the level of biological
organization under consideration as well as physiological state. Despite
this complexity, prominent single-cause models propose that metabolic
rates are supply-limited and that the scaling of supply systems provides
a sufficient explanation for the allometric scaling of metabolism. We
argue that some assumptions, as well as current interpretations of the
meaning (or consequences) of these constraints are flawed, i.e.,
elephants do not have lower mass-specific basal or maximal rates of
aerobic metabolism because their mitochondria are more supply-limited
than those of shrews. Animals do not violate the laws of physics, and
the allometric scaling of supply systems would be expected, to some
extent, to be matched by capacities for (and rates of) energy
expenditure. But life is not so simple. Animals are so diverse that to
do justice to metabolic scaling, it is also necessary to consider the
scaling of energy expenditure. It is by doing so that models of
metabolic scaling can be consistent with current paradigms in metabolic
regulation and accommodate the range of inter- and intraspecific
exponents found in nature. The "allometric cascade," a first attempt at
such an accounting, was a source of great satisfaction to Peter
Hochachka. It was the last door that he helped open to comparative
physiologists before he said goodbye
Czekala,
N.M., MacDonald, E.A., Steinman, K., Walker, S., Garrigues, N.W., Olson,
D., 2003. Estrogen and LH dynamics during the follicular phase of the
estrous cycle in the Asian elephant. Zoo Biology 22, 443-454.
Abstract: Pituitary and corpus luteum hormone patterns throughout the
elephant estrous cycle have been well characterized. By contrast,
analysis of follicular maturation by measurement of circulating
estrogens has been uninformative. This study tested the ability of a
urinary estradiol-3-glucuronide radioimmunoassay to noninvasively assess
follicular development during the nonluteal phase of the elephant
estrous cycle, and to determine the relationship between estrogen
production and the "double LH surge." Daily urine and serum samples were
collected throughout seven estrous cycles from three Asian elephants,
and urine was collected from an additional three females, for a total of
13 cycles. Serum was analyzed for luteinizing hormone (LH), and urine
was analyzed for estrogens and progestins. Elephants exhibited a typical
LH pattern, with an anovulatory LH (anLH) surge occurring approximately
21 days before the ovulatory LH (ovLH) surge. The urinary estrogen
pattern indicated the presence of two follicular waves during the
nonluteal phase. The first wave (anovulatory) began 5 days before the
anLH surge and reached a maximum concentration the day before the peak.
Thereafter, urinary estrogens declined to baseline for 2 weeks before
increasing again to peak concentrations on the day of the ovLH surge.
Urinary progestins were baseline throughout most of the follicular
phase, increasing 2-3 days before the ovLH surge and continuing into the
luteal phase. These results support previous ultrasound observations
that two waves of follicular growth occur during the nonluteal phase of
the elephant estrous cycle. Each wave is associated with an increase in
estrogen production that stimulates an LH surge. Thus, in contrast to
serum analyses, urinary estrogen monitoring appears to be a reliable
method for characterizing follicular activity in the elephant.
Davis, B.L.,
Dill, W.M., Hicks, A.R., Goodwin, T.E., Hollister-Smith, J., Alberts,
S.C. Use of SPME and GC-MS for chemical analysis of urine from African
elephants in musth. Abstracts Of Papers Of The American Chemical Society
225[614-CHED Part 1 MAR 2003]. 2003.
Ref Type: Abstract
Abstract: Addresses: Hendrix Coll, Dept Chem, Conway, AR 72032 USA ;Duke
Univ, Dept Biol Sci, Durham, NC 27706 USA
Dill, W.M.,
Davis, B.L., Hicks, A.R., Goodwin, T.E., Rasmussen, L.E.L., Loizi, H.,
Schulte, B.A. Chemical analysis of preovulatory female African elephant
urine: A search for putative pheromones. Abstracts Of Papers Of The
American Chemical Society 225[409-CHED Part I Mar 2003]. 2003.
Ref Type: Abstract
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.
Pucher, H.E.,
Stremme, C., Schwarzenberger, F., 2003. Priapism in a semiwild Asian
elephant (Elephas maximus) in Vietnam. Vet Rec 153, 717-718.
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.
Rasmussen,
L.E., Wittemyer, G., 2002. Chemosignalling of musth by individual wild
African elephants (Loxodonta africana): implications for conservation
and management. Proc R Soc Lond B Biol Sci 269, 853-860.
Abstract: Elephants have extraordinary olfactory receptive equipment,
yet this sensory system has been only minimally investigated in wild
elephants. We present an in-depth study of urinary chemical signals
emitted by individual, behaviourally characterized, wild male African
elephants, investigating whether these compounds were the same,
accentuated, or diminished in comparison with captive individuals.
Remarkably, most emitted chemicals were similar in captive and wild
elephants with an exception traced to drought-induced dietary cyanates
among wild males. We observed developmental changes predominated by the
transition from acids and esters emitted by young males to alcohols and
ketones released by older males. We determined that the ketones
(2-butanone, acetone and 2-pentanone, and 2-nonanone) were considerably
elevated during early musth, musth and late musth, respectively,
suggesting that males communicate their condition via these compounds.
The similarity to compounds released during musth by Asian male
elephants that evoke conspecific bioresponses suggests the existence of
species-free 'musth' signals. Our innovative techniques, which allow the
recognition of precise sexual and musth states of individual elephants,
can be helpful to managers of both wild and captive elephants. Such
sampling may allow the more accurate categorization of the social and
reproductive status of individual male elephants.
Sanchez,
C.R., Murray, S.Z., Montali, R.J., Spelman, L.H. Medical Management of
Acute Pylelonephritis in an Asian Elephant. Baer, C. K. American
Association of Zoo Veterinarians Annual Conference. 162-164. 2002.
2002.
Ref Type: Conference Proceeding
Singh, V.N.,
2002. Symptomatic study of haemorrhagic septicaemia in elephant in
Mudumalai Wildlife Sanctuary, Tamil Nadu. Indian Forester 128,
1089-1100.
Abstract: Symptoms of haemorrhagic septicaemia, a dreaded disease in
elephant is recorded in this study which reveals systematic spread of
Oedema from jowl to throat, neck, brisket, abdomen and perenical
regions. It also records the changes in character/colour of dung, urine,
eye, tongue, trunk, body temperature, feeding habit and body condition
along with treatment given to cure the disease.
Dehnhard, M.,
Hesitermann, M., Goritz, F., Hermes, R., Hildebrand, T., Haber, H.,
2001. Demonstration of 2-unsaturated C19-steroids in the urine of female
Asian elephants, Elephas maximus, and their dependence on ovarian
activity. Reproduction-Cambridge 121, 475-484.
Abstract: An oestrous-related pheromone of the female Asian elephant
(Elephas maximus) is known to induce behavioural responses in elephant
bulls. Additional data revealed that timing of oestrus in females with
close social relationships tends to be synchronized. Therefore, urine
from female Asian elephants might be expected to contain luteal
phase-dependent volatile substances, which may function as additional
chemical signals in this species. The aim of the present study was to
identify such compounds and to investigate their pattern of excretion
throughout the ovarian cycle. Urine samples were collected 3 times a
week during the follicular phase and 1 to 3 times a week during the
luteal phase from 5 adult female Asian elephants from a total of 13
non-conception cycles and one conception cycle, including the first 72
weeks of pregnancy. A simple headspace solid-phase microextraction
method has been developed for quantification of urinary volatile
substances and analysis was performed by gas chromatography. The
comparison of urine collected during the follicular and the luteal phase
indicated the presence of two luteal phase-dependent substances. Mass
spectrometry was used to identify one substance as
5alpha-androst-2-en-17-one and a second substance as the corresponding
alcoholic compound 5alpha-androst-2-en-17beta-ol. The
5alpha-androst-2-en-17beta-ol and -17-one profiles reflected cyclic
ovarian activity with clear (10-20-fold) luteal phase increases.
Furthermore, measurements of both compounds were correlated positively
with the concentration of urinary pregnanetriol and indicated cycle
duration (15.1±1.2 weeks) similar to that obtained from pregnanetriol
measurements (15.2±1.6 weeks). The results demonstrated the presence of
2 luteal phase-specific steroidal volatile compounds in elephant urine.
One of the substances, 5alpha-androst-2-en-17-one, has been demonstrated
in human axillary bacterial isolates. The measurement of both volatile
substances in elephant urine can be used for rapid detection of the
stage of the ovarian cycle, as the analysis can be completed within 2 h.
Ganswindt,
A., Heistermann, M., Borragan, S., Hodges, J.K., 2001. Assessment of
testicular endocrine function in captive African elephants by
measurement of urinary and fecal androgens. Zoo Biology 21,
27-36.
Rasmussen,
L.E.L., 2001. Source and cyclic release pattern of (Z)-7-dodecenyl
acetate, the pre-ovulatory pheromone of the female Asian elephant.
Chemical Senses 26, 611-623.
Abstract: Female Asian elephants (Elephas maximus) release a pre-ovulatory
urinary pheromone, (Z)-7-dodecenyl acetate (Z7-12:Ac), to signal males
of their readiness to mate. Z7-12:Ac is quantitatively elevated during
the follicular stage of oestrus, reaching maximum concentrations just
prior to ovulation, as demonstrated by two complementary headspace
techniques: (1) evacuated canister capture followed by cryogenic
trapping and (2) solid phase microextraction (SPME) used prior to gas
chromatography/mass spectrometry (GC/MS). These patterns were coincident
with observed male behaviours and were consistent with biochemical and
binding properties of the active ligand, including optimal binding pH.
To release maximum amounts of Z7-12:Ac for quantitation, serum and urine
samples from three mature female Asian elephants in their luteal and
follicular stages of several oestrous cycles were subjected to heat and
pH changes and were then treated with protease prior to SPME-GC/MS
analyses. When the post-luteal serum progesterone concentrations
declined to baseline levels, Z7-12:Ac became detectable in the female
urine. Throughout the follicular stage, pheromone concentrations
increased linearly with no apparent relationship to the two serum
luteinizing hormone peaks. Pre-ovulatory urine also contained related
compounds, including (Z)-7-12-dodecenol. The relative amount of this
alcohol increased relative to acetate during long-term storage, with a
proportional reduction in bioactivity. Z7-12:Ac was not detected in
mucus samples from the urogenital tract. A potential precursor of
Z7-12:Ac was identified in liver homogenates from female elephants in
the follicular stage. Erratum in: Chem Senses 2001 Sep;26(7):935
Rasmussen,
L.E.L., Krishnamurthy, V., 2001. Urinary, temporal gland and breath
odors from Asian elephants of Mudumalai National Park. Gajah 20,
1-7.
Schmid, J.,
Heistermann, M., Ganslosser, U., Hodges, J.K., 2001. Introduction of
foreign female Asian elephants (Elephas maximus) into an existing group:
behavioural reactions and changes in cortisol levels. Animal-Welfare 10,
357-372.
Abstract: The present study examined the extent to which the
introduction of three female Asian elephants (aged 3, 11 and 27 years)
into a group of 5 (1 male, 4 female) elephants at Munster zoo, Germany,
affects the behaviour and urinary cortisol levels of the animals
involved. At Munster, only the females were monitored , while the bull
was mainly kept separate. Behavioural observations were carried out
before transfer and during the six-month period following transfer.
Urine samples were collected regularly from each elephant during the
whole observation period. All the elephants showed behavioural changes
to the process of introduction. The transferred animals increased their
social behaviour after arrival in the foreign zoo. Two of them showed an
increase in stereotypies and one a reduction in stereotypes. The
elephants at Munster reacted with decreased frequencies of stereotypies
and increased frequencies of social behaviour and
manipulation/exploration behaviour. Six months after transfer, three of
the four elephants at Munster and one of the three transferred elephants
showed nearly the same behavioural activity pattern as before transfer.
One female still showed elevated stereotypic behaviour. From the four
elephants in which cortisol measurements could be reliably performed
(two of the transferred elephants and two elephants at Munster), only
one individual at Munster responded to the process of introduction with
a short-term elevation in urinary cortisol levels. One elephant showed a
negative correlation between locomotion and cortisol levels and one a
positive correlation between stereotypies and cortisol levels. Taken
together, the results suggest that transfer and introduction caused some
stress responses in the elephants, but that stress was neither prolonged
nor severe. Serious welfare problems may have been prevented through
individual behavioural coping mechanisms and former experience with
stressful situations.
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.
Bielecki, W.,
Sobczak-Filipiak, M., Pietrak, M., Goslawski, J., Blaszczak, B., 2000. A
case of nephritis in an elephant. Zycie-Weterynaryjne 75,
197-199.
Lazar, J.,
Prestwich, G.D., Rasmussen, L.E.L., 2000. Urinary and trunk mucus
protein carriers of (Z)-dodecenyl acetate, the sex pheromone of the
Asian elephant. Chemical Senses 25, 603.
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.
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.
1999. Equine
Medicine and Surgery. Mosby, St. Louis MO USA.
Fies, M.,
Heistermann, M., Hodges, J.K., 1999. Patterns of urinary and fecal
steroid excretion during the ovarian cycle and pregnancy in the African
elephant (Loxodonta africana). Gen Comp Endocrinol 115, 76-89.
Abstract: The aims of the present study were to (I) determine the
relative abundance of the 5alpha-reduced progestins
5alpha-pregnane-3-ol-20-one (5alpha-P-3OH) and
5alpha-dihydroprogesterone (5alpha-DHP) and progesterone (P4) in African
elephant feces and to establish improved fecal progestin assays for
monitoring ovarian function; and (ii) describe longitudinal profiles of
urinary and fecal progestin and estrogen metabolites during pregnancy.
Matched urine and fecal samples were collected weekly from six adult
females throughout 18 nonfertile cycles and two complete pregnancies (89
and 93 weeks duration). Fecal samples were lyophilized and extracted
with 80% methanol in water and immunoreactive 5alpha-P-3OH, 5alpha-DHP,
and P4 and (for pregnant females only) estrone (E1) and estradiol (E2)
determined by enzyme immunoassay. Urine samples were hydrolyzed,
ether-extracted, and assayed for 5alpha-P-3OH, E1, and E2. HPLC
cochromatography of fecal extracts with various radioactive progestin
tracers confirmed the presence of large amounts of both 5-reduced
progestins (5alpha-P-3OH > 5alpha-DHP) but not of P4. 5-Reduced
progestins (but not P4) were excreted in a cyclic pattern and levels
were significantly correlated with urinary 5alpha-P-3OH. Fecal
5alpha-P-3OH showed the more pronounced and consistent luteal-phase
elevation and a better correspondence to urine with respect to timing of
the luteal-phase rise. Fecal and urinary 5-reduced progestins increased
gradually during early pregnancy to maximum values around week 40-45.
Levels gradually declined during the second half of pregnancy, reaching
baseline values 2 days before parturition. Urinary estrogens did not
show any cyclic pattern during the preconception period and levels
remained low during the first 30 weeks of gestation. Thereafter, there
was a rapid 10- to 20-fold increase to maximum values at mid-pregnancy,
followed by a gradual decline to birth. There was no mid-pregnancy
elevation in fecal estrogens, but there was a modest increase in E1
during the second half of gestation.
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.
Gual-Sill,
F., Pickard, A.R., Holt, W.V., Green, D. Preliminary Results of
Non-Invasive Monitoring of the Estrous Cycle in Female Asian Elephants (Elephas
maximus) through Fecal Steroid Analysis. 1999 Proceedings American
Association of Zoo Veterinarians. 87-92. 1999. 1999.
Ref Type: Conference Proceeding
Abstract: For a number of years, estrous cycle monitoring and pregnancy
detection in the Asian elephant has been performed using urinary steroid
hormone metabolite analysis; this technique presents some practical
problems. Monitoring the reproductive status through fecal steroid
analysis is possible in this and many other species. The steroid
metabolite profiles of female Asian elephants were monitored by
enzyme-linked immunosorbent assay (ELISA), to provide detailed
information about the estrous cycle and pregnancy in this species, and
to investigate causes of reproductive failure. Fecal and matched urine
samples were non-invasively collected regularly for 6 mo from captive
female Asian elephants. (n = 4 cyclic; n = 1 acyclic). The samples were
frozen at -20ºC. Gas Chromatography and Mass Spectroscopy (GC-MS)
procedures were used to investigate the steroid hormone metabolite
profile and to identify the major excretory metabolites; no steroid
metabolites were found in the concentrated extracted feces of this
species using the currently available methodology. The fecal
pregnanetriol profile observed in three of the cyclic females showed a
clear relation with their matched urinary pregnanetriol profile and a
cyclic pattern was demonstrated. Fecal pregnanetriol values increased
from an overall mean of 94.67 ng/g of dry feces (+/- 13.24, range 31.5 -
219.12 ng/g) during the inter-luteal period to a luteal phase mean of
334.61 ng/g dry feces (+/- 43.48, range 34.35 - 1035.1 ng/g). All the
data collected from the fecal and urinary analysis of pregnanetriol in
all five individuals investigated demonstrated a significant
relationship between urinary and fecal pregnanetriol. The acyclic
individual showed a mean fecal pregnanetriol concentration of 84.91 ng/g
(+/- 13.06) and values ranged from 33.17 ng/g to 211.42 ng/g. Fecal
steroid hormone metabolite analysis for monitoring estrous cycles in
Asian elephants may be used in the future to monitor free-roaming, wild
or semi-wild individuals as well as those in captivity to assist
reproductive and conservation programs of this highly endangered
species.
Rasmussen,
L.E.L., 1999. Evolution of chemical signals in the Asian elephant,
Elephas maximus: behavioural and ecological influences. Journal of
Biosciences 24, 241-251.
Abstract: In antiquity, the Asian elephant, Elephas maximus,
gradually spread southward and eastward to become a successfully
surviving, ecologically dominant megaherbivore in the tropical
environment of south-east Asia. The changing physical environment forced
dynamic fluxes in its social structure and altered its metabolism. Such
events shaped the production and ultimately the stability of certain
chemicals released by body effluvia. Some of these chemicals took on
significance as chemical signals and/or pheromones. This article
demonstrates by experimental and observational evidence, and
hypothesizes based on speculative reasoning, how and why specific
chemical signals evolved in the modern Asian elephant. Evidence,
including the functional criteria required by elephant social structure
and ecology, is presented for the hypothesis that the recently
identified female-emitted, male-received sex pheromone, (Z)-7-dodecenyl
acetate evolved first as a chemical signal. Subsequently, the
cohesiveness and harmony of small, matriarchally-led female groups were
strengthened by a female-to-female chemical signal, recently defined
behaviourally. The looser societal structure of freer, roaming males
also became bounded by chemical signals; for the males, breath and
temporal gland emissions, as well as urinary ones function in chemical
signaling. Basic knowledge about elephant chemical signals is now
linking chemical information to behaviour and beginning to demonstrate
how these signals affect elephant social structure and enable the
species to cope with environmental changes.
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.
Rasmussen,
L.E.L., Schulte, B.A., 1999. Ecological and biochemical constraints on
pheromonal signaling systems in Asian elephants and their evolutionary
implications. In: Johnston, R.E., Muller-Schwarze, D., Sorenson, P.W.
(Eds.), Advances in Chemical Communication in Vertebrates 8. Kluwer/Academic/
Plenum Press, pp. 49-62.
Abstract: The Asian elephant is an unusual example of how intraspecies
chemical communication helps maintain societal cohesiveness within
familial and herd units. The amount of multi-directional chemical
communication is surprising, because long-lived elephants have a highly
organized society, are capable of trans-generational passage of
information, possess a sophisticated vocalization system, and are
capable of complex learning and tool use. This paper discusses the
ecological, behavioral, and biochemical aspects of chemical signals in
elephants from an evolutionary perspective. Diverse bodily emissions are
utilized as intraspecies chemical signals (including pheromones), often
with imposed biochemical constraints. In this chapter, chemosignals
released from the temporal gland secretions and breath of male Asian
elephants in musth and a urinary female-to-male preovulatory pheromone
are utilized as examples of these concepts. Furthermore, specific
behavioral and biochemical studies with (Z)-7-dodecenyl acetate (a
female-to-male urinary sex pheromone) demonstrate that social context
significantly influences responsivity (demonstrated by field studies in
Myanmar) and that additional biochemical requirements, perhaps lipocalin-like
proteins, may be required for full bioactivity. The remarkable
convergent evolution of (Z)-7-dodecenyl acetate, both structurally and
functionally, in elephants and Lepidoptera, allows the use in elephant
studies of effective biochemical tools developed for insect
investigations. This convergence of chemical signaling systems of
elephants and insects has several interesting implications.
Schulte,
B.A., Rasmussen, L.E., 1999. Signal-receiver interplay in the
communication of male condition by Asian elephants. Animal Behaviour 57,
1265-1274.
Abstract: Signal design and meaning are dependent on the condition of
the sender and receiver as well as the response of the receiver. This
study examined (1) whether female Asian elephants, Elephas maximus, can
distinguish between a conspecific male in musth and nonmusth states
using urinary signals, (2) how the oestrous condition of the female
affects discrimination, and (3) correlation of female responses with the
testosterone level of the male. Musth is a rut-like state displayed by
healthy adult male elephants. Males in musth dominate nonmusth males and
may be preferred by females as mates. Urine was collected from two
captive male Asian elephants during nonmusth periods and from one of
these males during times of musth. Samples of musth and nonmusth urine
and control liquids were placed in an elephant enclosure weekly for 16
weeks, the length of a female oestrous cycle. Primary response
behaviours were approach and four trunk-tip motions, namely sniff,
check, place and flehmen. Musth urine consistently elicited greater
responses than nonmusth and control samples. Females were more
responsive during their follicular (sexually receptive) than luteal
(unreceptive) stages of oestrus. Furthermore, females appeared to be
sensitive to the degree of musth as responses increased with rising
serum testosterone levels of the male donor. Chemical signals from males
are a likely source of honest signals related to status and reproductive
condition. Female elephants appear capable of detecting differences in a
male based upon urinary chemosignals.
Wrench, A.,
Kock, N.D., Gordon, S.J.G., Turner, A.D., Fries, D.S., 1999.
Investigation of selected plants as possible causes of flaccid trunk
paralysis in free-ranging elephants (Loxodonta africana) in Zimbabwe.
Zimbabwe Veterinary Journal 30, 93-97.
Abstract: Extracts from four plants collected from the southern shore of
Lake Kariba were injected into Balb/c laboratory mice in an attempt to
establish the cause of flaccid paralysis of the trunk in elephants.
Behavioural and neurological changes were noted in some instances after
injection. Histopathological changes similar to those found in affected
elephants, including renal tubular karyomegaly and myocardial
degeneration and necrosis, were found in mice exposed to Boerhavia
diffusa. Patchy gliosis and cerebral oedema were present in mice treated
with Blumea geriepina and Geigeria schnizii. The study indicates the
need for further investigation into the role of plant intoxication as a
possible cause of trunk paralysis in elephants.
Hodges, J.K.,
1998. Endocrinology of the ovarian cycle and pregnancy in the Asian
(Elephas maximus) and African (Loxodonta africana) elephant. Animal
Reproduction Science 53, 3-18.
Abstract: In reviewing the literature, this paper assesses the current
level of understanding of the hormonal control of the ovarian cycle and
pregnancy in the
elephant. Data are compiled from two genera, Elephas maximus and
Loxodonta africana, recognizing differences where known to exist.
Measurements of various steroid and peptide hormones, but most notably
immunoreactive progesterone (iP), indicate an overall cycle length of
15-16 weeks, comprising an 8-11-week luteal phase and a shorter inter-luteal
(follicular) period of 4-6 weeks. Oestrogen related phenomena suggest
the occurrence of two (possibly more) waves of follicular development
with a 3-week periodicity, although oestrogen levels are low and
fluctuate without clear pattern. The inter-luteal period is
characterized by two discrete LH peaks also 3 weeks apart, but only the
second is associated with an iP rise. Ovulation rate is variable and
additional luteal structures lacking ovulation points are probably
formed each cycle. The post-ovulatory period is associated with elevated
progestogen levels, mainly 5alpha-reduced compounds, while progesterone
itself is a minor secretory product. A revised model for the elephant
ovarian cycle is presented. Detailed information on the endocrinology of
pregnancy is lacking. Elevated progestins beyond the normal luteal phase
indicate that conception extends the functional lifespan of the CL, with
maximum steroidogenic activity between 3-15 months. The nature of the
luteotrophic support is not known and placental gonadotrophins have not
been conclusively demonstrated. Progestins fluctuate at or slightly
above luteal phase values throughout pregnancy. There is a marked
increase in prolactin levels after 16-24 weeks and a relationship with
oestrogen secretion may exist since circulating and urinary levels show
a progressive increase from 6-8 months. A placental contribution to
progestin secretion is likely, although direct evidence is lacking.
Considerable enlargement of foetal gonads during the second half of
pregnancy in African elephants suggests a possible steroidogenic
function, but none has been defined. The endocrine events leading to
parturition are unknown. In the Asian elephant, a fall in iP precedes
parturition; oestrogen levels decline and cortisol increases at the time
of birth. The paper concludes with a brief selection of priorities for
future research.
Rasmussen,
B., 1998. The Chemical Identification of a Preovulatory Pheromone: A
Reproductive Chemosignal from Female to Male Asian Elephants. Journal of
the Elephant Managers Association 7, 52-56.
1997.
Laboratory Procedures for Veterinary Technicians. Mosby, St. Louis MO.
Heisterman,
M., Trohorsch, B., Hodges, J.K., 1997. Assessment of ovarian function in
the African elephant (Loxodonta africana) by measurement of
5-reduced progesterone metabolites in serum and urine. Zoo
Biology 16, 273-284.
Abstract: We have previously shown that 5-pregnane-3,20-dione (5-DHP)
and 5-pregnane-3-ol-20-one (5-OH) are the major luteal and circulating
progestins in the African elephant. Therefore, the aim of the present
study was to determine 1) circulating levels and patterns of secretion
of 5-DHP in relation to progesterone (P4) throughout the ovarian cycle,
2) the presence and relative abundance of 5-reduced progestins in urine
and 3) whether their measurement in urine would provide a non-invasive
method for monitoring luteal function. Urine samples were collected
weekly throughout a total of 13 ovarian cycles from 5 females. In
addition, matched blood samples were collected during 6 cycles from 2 of
the 5 animals. All hormone measurements were carried out by
enzymeimmunoassay following extraction. Urine was hydrolized prior to
extraction. Profiles of P4 and 5-DHP in serum followed a similar
cyclic pattern and both measurements were significantly correlated (r =
0.78-0.98, mean 0.89, P < 0.001). Concentrations of 5-DHP were,
however 10-20-fold higher than those if P4. In addition, 5-DHP
measurements showed a more pronounced luteal phase increase than that of
immunoreactive P4. HPLC co-chromatography confirmed the presence of
large amounts of 5-P-3-OH in urine as a single immunoreactive peak,
whereas 5-DHP was present in very low levels and measurable only as one
of several immunoreactive substances. Measurements of urinary 5-P-3-OH
were significantly correlated to serum 5 -DHP measurements in each of
the 6 cycles (r = 0.72-0.93, mean 0.81, P < 0.001), whereas
correlation coefficients between urinary and serum 5-DHP values were
generally lower (r = 0.34-0.83, mean 0.69) and significant in only 4 of
6 cycles. Accordingly, only urinary excretion of 5-P-3-OH, but not of
5-DHP, exhibited a clear cyclic pattern, with consistently low levels
of 0.15-.020 g/mg Cr in the follicular phase and 10-fold elevated
levels (1.8-2.2 g/mg Cr) in the luteal phase. Based on the intervals
between successive luteal phase increases in urinary 5-P-3-OH, a mean
cycle length of 14.1+/- 1.8 weeks, comprising a follicular phase of 5.0
+/- 0.9 weeks and a luteal phase of 9.10+/-01.4 weeks was determined for
the 13 cycles studied. The results indicate that measurements of
5-P-3-OH in urine provide a reliable non-invasive method for monitoring
luteal function in the African elephant.
Niemuller,
C., Shaw, H.J., Hodges, J.K., 1997. Pregnancy determination in the Asian
elephant (Elephas maximus): A change in the plasma progesterone
to 17 hydroxyprogesterone ratio. Zoo Biology 16, 415-426.
Abstract: The measurement of circulating progesterone (P) is widely used
to confirm and monitor pregnancy in the captive Asian elephant.
Consistently elevated progesterone (P) concentrations for a minimum of
12 weeks can be used as a positive indication of pregnancy, although
two instances of prolonged P secretion during the cycle have been
recorded [Rubel, 1987; Olsen et al, 1994]. Previously, we demonstrated
that pregnanetriol was the major urinary gestagen metabolite enabling
the noninvasive monitoring of the reproductive cycle of the Asian
elephant [Niemuller et al., 1993] as well as pregnancy [unpublished
data]. the importance of this unusual urinary metabolite triggered an
investigation into the secretion of circulating concentrations of 17
hydroxyprogesterone (17 OHP) during pregnancy and reproductive cycles,
as this steroid is the only steroid precursor of pregnanetriol.
Comparison of the profiles between 17 OHP and P during early pregnancy
(n = 5) and nonconceptive cycles (n = 15) demonstrated a decline in 17
OHP , but not P, as early as week 3 postmating (designated as week 1)
and lasting up to week 13. Otherwise, secretions of 17 OHP mimicked P
concentrations throughout pregnancy and in nonconceptive cycles.
Examination of the mean ratio values of 17 OHP to P demonstrated a
significant drop in the ratio during weeks 2-7 of early pregnancy from
>/= 0.7 to < 0.7 compared with the same time period in a nonreceptive
cycles (p < 0.05m N = 5), A 2x2 table analysis of the 17 OHP:P ratio
during weeks 2-7 indicated that the possibility of a false positive or
false negative result was 3.4 and 6.5%, respectively, based on the
sensitivity and specificity of the test. Overall, gestation lengths of
the pregnancies completed during this study (N = 4) ranged from 91 to 98
weeks, with a mean of 93+/-2.9 (SEM) weeks. A birth-to-conception
interval of 47 weeks was noted in one animal. The results described in
this report provide additional data on the reproductive endocrinology of
the pregnant Asian elephant and also present the earliest means to date
of determining pregnancy by analysis of the 17 OHP:P ratio.
Kirkpatrick,
J.F., Turner, J.W., Jr., Liu, I.K., Fayrer-Hosken, R., 1996.
Applications of pig zona pellucida immunocontraception to wildlife
fertility
control. J Reprod Fertil Suppl 50, 183-189.
Abstract: A unique application of pig zona pellucida (PZP)
immunocontraception is the control of wildlife populations. A native PZP
vaccine has been successfully applied to wild horse and donkey
populations. A single annual booster inoculation was capable of
maintaining contraception. Seven consecutive years of PZP treatment in
wild mares resulted in no detectable debilitating side effects, and
reversibility of contraception has been documented among mares treated
for up to 4 consecutive years. Long-term treatment (5-7 years) is
associated with some ovulation failure and depressed urinary oestrogen
concentrations. Complex social behaviours in horses were unaffected by
treatment. PZP immunocontraception has also been successfully applied to
white-tailed deer, with no detectable changes in ovarian histology after
2 years of treatment. Seventy-four species of captive zoo animals have
been treated with the PZP vaccine, with documented success in 27
species, including members of the orders
Perissodactyla (Equidae), Artiodactyla (Cervidae, Capridae, Giraffidae,
Bovidae), and Carnivora (Ursidae, Mustelidae, Felidae).
Immunocytochemistry studies have demonstrated a high degree of
crossreactivity between anti-PZP antibodies and African elephant zona
pellucida. The need for a one-inoculation form of the vaccine has led to
the incorporation of PZP into lactide-glycolide microspheres, which
cause a delayed release of the PZP. PZP immunocontraception of wildlife
has potential because of (1) > 90% effectiveness, (2) the ability for
remote delivery, via darts, (3) reversibility after short-term use, (4)
a wide breadth of effectiveness across many species, (5) a lack of
debilitating side-effects even after long-term treatment, and (6)
minimal effects upon social behaviours.
Murray, S.,
Bush, M., Tell, L.A., 1996. Medical management of postpartum problems in
an Asian elephant (Elephas maximus) cow and calf. Journal of Zoo
and Wildlife Medicine 27, 255-258.
Abstract: An 18-yr old female Asian elephant (Elephas maximus)
gave birth to a 120-kg female calf following 22 mo of gestation.
Immediately after parturition, the cow became agitated and aggressive
towards the calf. Before the keepers were able to safely intervene and
remove the calf, the cow stepped on the calf's head and right front
leg. Within 30 min, the cow calmed down, allowing the calf's safe
reintroduction under close keeper supervision and control. The cow had
a retained placenta, poor mammary development, and low milk production.
The calf's injuries, in combination with the cow's low milk production,
impeded the calf's ability to nurse and gain weight. Within 10 days,
the calf lost 10% of its weight. Serum protein electrophoresis
indicated failure of passive transfer of maternal immunoglobulin. On
day 10, the calf received a transfusion of concentrated immunoglobulin
extracted and concentrated from the cow's previously banked plasma. On
day 13, the calf developed a urinary tract infection, as diagnosed by
white blood cells and bacteria in the urine. Following immunoglobulin
administration and antibiotic therapy, clinical signs slowly resolved
and the calf gained weight. The cow passed the fetal membranes during
parturition, but the placenta was retained. Despite prophylactic
systemic antibiotics and vaginal flushing, the cow became depressed and
developed a leukocytosis and anemia. A mucopurulent vaginal discharge
and ventral edema were noted on day 3, and milk production was minimal.
Because decreased milk production has been reported as a common sequel
to retained placenta, efforts were focused on removing the placenta.
Intermittent oxytocin therapy on days 2-14 did not result in expulsion
of the placenta and produced only transient abdominal contractions and
minimal increases in milk letdown. On day 15, 10 mg estradiol cypionate
was administered i.m. followed by 200 IU oxytocin i.v. An additional 10
IU oxytocin was administered i.v. on day 16. The friable placenta was
palpable within the vaginal vault on day 17. The remaining placenta was
removed by gentle traction applied by a modified weighted pressure
cuff. Once the placenta was removed, the cow's clinical problems slowly
resolved and the calf continued to gain weight.
Rasmussen,
L.E.L., Lee, T.D., Roelofs, W.L., Zhang, A., Daves, G.D., 1996. Insect
pheromone in elephants. Nature 379, 1.
Brown, J.L.,
Wemmer, C.M., Lehnhardt, J., 1995. Urinary Cortisol Analysis for
Monitoring Adrenal Activity in Elephants. Zoo Biology 14 ,
533-542.
Abstract: Cortisol was measured in dichloromethane-extracted elephant
urine using an 125I solid-phase radioimmunoassay (RIA). The
cortisol RIA was validated by demonstrating 1) parallelism between
dilutions of pooled urinary extracts and the standard curve, 2)
significant recovery of exogenous cortisol added to elephant urine, and
3) a relationship between changes in the peripheral and urinary cortisol
after an adrenocorticotropin hormone (ACTH) challenge. One African (Loxodonta
africana) and one Asian (Elephas maximus) elephant were given
three injections of ACTH (1.25 mg) at 2 h intervals. Serum cortisol
increased four- to eightfold within 30 min after the first injection and
peaked (nine- to twelvefold increase) after the second injection. Serum
concentrations began to decline 2-3 h after the last injection but were
still approximately fourfold higher than baseline at the end of the
collection period (hour 8). In the urine, cortisol concentrations were
increased in the first sample postinjection (1.5 - 4 h) and peaked
twenty- to fortyfold by ~6 h. Urinary cortisol remained elevated at 8 h,
but returned to baseline by the following morning. Analysis of high
performance liquid chromatography fractions of extracted urine revealed
that immunoactivity was associated with free cortisol (~90% of total
immunoactivity) and a more polar, unidentified metabolite. A method for
preserving urine was developed to allow storing unfrozen samples. One
pool of urine from each of one African and two Asian elephants was
divided into aliquots, placed in tubes containing absolute ethanol
(10%), sodium azide (0.1%) or distilled water (control), and frozen
after 0, 1 , 2, 3, 4, 6, 8, 10, 12, and 24 weeks of storage at ~25ºC.
In unpreserved samples, cortisol concentrations were reduced 46% by 2
weeks and 95% by 24 weeks. In contrast, ethanol- and sodium azide-preserved
samples retained 100 and 95% of cortisol immunoactivity through 8 weeks
and 93 and 85% of activity through 12 weeks, respectively. We infer from
these data that changes in urinary cortisol excretion in the elephant
reflect fluctuations in adrenal activity and may be a useful indicator
of stress. Additionally, urine samples can be collected and stored
unfrozen for at least 2 months before any appreciable loss in cortisol
immunoactivity occurs, a finding potentially useful to field application
of this technique.
Brown, J.L.,
Lehnhardt, J., 1995. Serum and urinary hormones during pregnancy and the
peri- and postpartum period in an Asian elephant (Elephas maximus).
Zoo Biology 14, 555-564.
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.
1994.
Veterinary Laboratory Medicine. Iowa State University Press, Ames, IA.
Lodwick, L.J.,
Dubach, J.M., Phillips, L.G., Brown, C.S., Jandreski, M.A., 1994.
Pharmacokinetics of amikacin in African elephants (Loxodonta africana).
Journal of Zoo and Wildlife Medicine 25, 367-375.
Abstract: Two adult females were given single i.v. injections of 8
mg/kg. Trials using 3 mg/kg and 6 mg/kg i.m. were conducted with three
adult females. Serum concentrations of amikacin were measured serially
over a 24-49 h period. After i.v. administration of 8 mg/kg, the
elimination half-lives (t0.5) were 4.0 and 3.7 h, the volumes of
distribution at steady state were 0.21 and 0.18 litres/kg, and total
body clearances were 41.8 and 40.8 ml/h/kg. At i.m. doses of 3 and 6
mg/kg, the peak concentrations observed ranged from 4.8 to 8.4 µg/ml and
14.2 to 21.8 µg/ml, respectively. The time at observed peak
concentration was between 1 and 3 h, and t0.5 ranged from 3.8 to 5.9 h
for the lower dose and from 3.7 to 6.3 h for the higher dose. Following
the single dose trials, one elephant was treated with amikacin at a dose
of 7 mg/kg i.m. at 24 h intervals for 21 days, and serum amikacin
concentrations were determined 2 to 4 times on each of 11 days. Mean
(SD) peak serum concentration for this elephant was 19.0±2.8 µg/ml, and
mean serum concentration at 24 h (through) was 1.7±0.4 µg/ml. There was
indication in this one elephant of a mild, reversible renal tubular
insult based on a slight transient elevation in serum creatinine and the
presence of casts in the urine. These changes resolved soon after the
end of treatment. These preliminary results suggest that amikacin
administered at 6-8 mg/kg i.m. once every 24 h would be appropriate for
elephants with bacterial infections suspected to be susceptible to
amikacin.
Plouzeau, E.,
daCunha, S., Shaw, H.J., 1994. The ovarian cycle in Asian and African
elephants (Elephas maximus and Loxodonta africana). Techniques for the
monitoring of female fertility in captivity. Revue de Medecine
Veterinaire 145, 905-911.
Abstract: A discussion. During the oestrous cycle, circulating blood
progesterone in elephants alternates between high and low concentrations
over a cycle of 16±2 weeks. Some of the data have suggested a 3-week
cycle and other data a 15- to 16-week cycle. Captive females show no
signs of oestrus, although males show a Flehmen-like response to female
urine, which is inversely related to plasma progesterone concentration.
Pregnancy is associated with a sustained increase in circulating
progesterone concentration and with an increase in total oestrogen,
prolactin and oestrogen concentrations during the second half of
pregnancy. The most reliable method of monitoring the oestrous cycle and
pregnancy is by weekly analysis of plasma progesterone in both species,
or of its urinary metabolite (pregnanetriol) in Asian elephants.
Wallace, C.,
Byron, T.H., Foerner, J.J., Weston, H., Kilpatrick, J., Jastremski, M.S.
Clinical case report: the medical management and treatment of a 36 year
old premiparturient Asian elephant cow with a dystocia and following a
Caesarian section. 1994.
Ref Type: Unpublished Work
Abstract: The medical history and management of a 36 year old
premiparturient Asian elephant (Elephas maximus indicus) with a
dystocia requiring a caesarian section are discussed. The examination
and complete medical evaluation to determine the health of the cow and
viability and position of the calf are described. The medical
management of the post-operative complications and the changes in
complete blood counts, differential, serum biochemistry values and
urinalysis prior to the elephant's death are described. Complications
included peritonitis with systemic sepsis, renal failure, and hepatic
failure. Pertinent necropsy findings on the cow included severe diffuse
subacute peritonitis, uterine transmural necrosis, diffuse renal tubular
nephrosis, and hepatic centrolobular degeneration.
Niemuller,
C.A., Shaw, H.J., Hodges, J.K., 1993. Non-invasive monitoring of ovarian
function in Asian elephants (Elephas maximus) by measurement of urinary
5beta-pregnanetriol. Journal of Reproduction and Fertility 99,
617-625.
Abstract: An enzymeimmunoassay for 5beta-pregnanetriol is described.
Immunoreactive pregnanetriol concentrations were significantly
correlated with the concentrations of progesterone (0.98, n = 269,
P<0.01) and 17alpha-hydroxyprogesterone (0.95, n = 205, P<0.01), the
precursor of pregnanetriol. The duration of cycles as determined by
measurements of plasma progesterone, plasma 17alpha-hydroxyprogesterone
and urinary pregnanetriol averaged 15.54±1.5 weeks (23 cycles),
15.21±1.7 weeks (15 cycles) and 15.45±0.94 weeks (20 cycles)
respectively. The results demonstrated that it is possible to monitor
ovarian function in Asian elephants by measuring urinary pregnanetriol
concentration.
Okewole,
P.A., Oyetunde, I.L., Irokanulo, E.A., Chima, J.C., Nwankpa, N., Laleye,
Y., Bot, C., 1993. Anthrax and cowdriosis in an African elephant
(Loxodonta africana). Veterinary Record 133, 168.
Abstract: In February 1992, a 15-year-old African elephant died; it was
the second elephant that had died within 2 weeks at a wildlife park.
Clinical signs in both elephants included frequent micturition,
restlessness and weakness of the hindquarters with frequent falls. PM
examination revealed ecchymosis of the epicardium, atrioventricular
surfaces of the heart and serosal surfaces of the intestines and bladder
with sloughing of intestinal mucosae. The liver was enlarged, ecchymotic
and congested. A serosanguinous exudate with fibrin was present in the
thoracic and abdominal cavities. The meninges were congested. Bacillus
anthracis was cultured from tissue samples and from tissue samples from
guineapigs inoculated with broth cultures of the tissue samples from the
elephant. Cowdria ruminantium was identified in stained impression
smears from the elephant brain. This appears to be the first report of
the simultaneous occurrence of anthrax and cowdriosis in an African
elephant.
Turczynski,
C.J. The endocrinology of musth in the male Asiatic elephant (Elephas
maximus): Serum estradiol, serum LH and serum, fecal and urinary
testosterone. 1993. College Station, TX. USA, Texas A&M University.
Ref Type: Thesis/Dissertation
Czekala,
N.M., Roocroft, A., Bates, M., Allen, J., Lasley, B.L., 1992. Estrogen
metabolism in the Asian elephant (Elephas maximus). Zoo Biology
11, 75-80.
Abstract: Estradiol-17B metabolism was studied in two female Asian
elephants (Elephas maximus). In an initial study, 500 microCi of
tritiated estradiol-17B was injected iv into a single animal, and 0, 30,
and 60 min serum samples were collected as well as all excreted urine
and feces for 24hr. In a second study, 1.5 mg unlabeled estradiol-17B
was injected iv into a second animal and 0, 5, 15, 30 and 60 min serum
samples and a 30 min urine sample were collected postinjection. Analyses
of samples from both studies demonstrated a rapid conversion of free
estradiol to conjugated forms in the serum. The first (5 min) serum
sample following the injection of unlabelled estradiol contained
unconjugated estradiol:conjugated estradiol: conjugated estrone at a
ratio of 60:29:10, respectively, and at 30 min a ratio of 33:43:24. The
urinary estrogen metabolites were in the conjugated form with an
estradiol:estrone ratio of 60:40. No radiolabelled estrogen was found
in the fecal samples during the 24 hr following administration of the
radiolabelled estradiol. These data indicate a rapid clearance of
circulating free estradiol in the elephant, with a major metabolite in
the serum and urine being estradiol conjugate.
Hagenbeck, D.
Attempts to monitor oestrus and pregnancy in Indian elephants by urine
testing. Diagnostische Zyklusansprache und Trachtigkeitsbestimmung beim
asiatischen Elefanten (Elephas maximus). 1-66. 1992. Hannover, Germany,
Tierarztliche Hochschule Hannover.
Ref Type: Thesis/Dissertation
Chakraborty,
A., Islam, S., Gogoi, A.R., Chaudhury, B., 1991. A note on clinical
examination of elephants in Manas Tiger Project and Kaziranga National
Park in Assam. Zoos' Print Journal November.
Abstract:
The diseases of elephants have been recorded by Steel (1885) and Evans
(1910) and till then many works have done on elephants. Compared to the
extent of work done in other states of our country, very little
investigation seems to have been done in Assam. The present
communication relates the report of investigation carried out in the
domesticated elephants of Manas Tiger Project and Kaziranga National
Park in Assam.
1990. The
story of Babe, the Asian elephant. Veterinary Viewpoints 2.
Diaz-Samayoa-de-Aguirre,
L. Sex hormones in blood plasma, urine and faeces from female and male
Indian elephants. Vergleichende Hormonuntersuchungen in Blutplasma, Harn
und Kot beim weiblichen und mannlichen asiatischen Elefanten (Elephas
maximus). 1-145. 1990. Hannover, Germany, Tierarztliche Hochschule.
Ref Type: Thesis/Dissertation
Mainka, S.A.,
Lothrop, C.D., Jr., 1990. Reproductive and hormonal changes during the
estrous cycle and pregnancy in Asian elephants. Zoo Biology 9,
411-419.
Abstract: Serum progesterone and urinary total estrogen concentrations
were determined weekly to bi-weekly in 2 female Asian elephants for 96
weeks. The mean estrous interval was approximately 16 weeks in the
nonpregnant animal. A total of 5 cycles were observed in the 96 weeks
study period. The serum progesterone concentration ranged from 150
pg/ml to greater than 350 pg/ml during the luteal phase of the estrous
cycle. The serum progesterone was elevated for 8-12 weeks of the 16 week
estrous cycle. The urinary total estrogen concentration ranged from
less than 10 to greater than 300 pg/microgram creatinine. The second
animal was pregnant at the beginning of the study period. The serum
progesterone concentration was elevated (> 100 pg/ml) in the pregnant
animal until parturition. The urinary total estrogens increased from
approximately 50 pg/microgram creatinine to greater than 400
pg/microgram creatinine during the first year of pregnancy and remained
elevated until parturition. Estrous cycling had not resumed by 3 months
post partum.
Brannian,
J.D., Griffin, F., Terranova, P.F., 1989. Urinary androstenedione and
luteinizing hormone concentrations during musth in a mature African
elephant. Zoo Biology 8, 165-170.
Abstract: Musth has not been well documented in captive African
elephants. A 37-year-old African bull elephant in the Kansas City
Zoological Park was observed during periods of behavioral musth and non-musth.
Androstenedione and luteinizing hormone (LH) concentrations in urine
were measured by radioimmunoassay. Urinary androstenedione and LH levels
were significantly higher in musth urine than in non-musth samples. A
positive correlation (P> 0.001) existed between urinary LH and
androstenedione concentrations. These results indicate that musth can
occur in a zoo-maintained African elephant and that urinary androgen
levels are elevated during musth, possibly as a result of LH stimulation
of testicular steroidogenesis.
Czekala,
N.M., Roocroft, A., Bates, M. Estrogen metabolism in the Asian elephant
(Elephas maximus). Biology of Reproduction 40[suppl.1], 119.
1989.
Ref Type: Abstract
Abstract: Understanding follicular development in elephants has been
confounded by an inability to detect serum estradiol (E2) in
cyclic patterns. Serum levels remain < 20 pg/ml throughout the cycle.
To understand E2 dynamics in the elephant, two metabolic
studies were initiated. First 500 microCi tritiated E2 was
injected (iv). All urine and feces voided during the following 24 hrs.
and two blood samples (30 min and 2 hrs) were collected. No radiolabel
appeared in feces during the 24 hrs. Urinary radiolabel analyzed by
HPLC appeared 100% conjugated, 52% E2-3-conjugate (E2C)
and 48% estrone conjugate (E1C). Peak levels appeared in
urine 30 min post-injection, decreased rapidly during the next 2 hrs,
then gradually to 16.5 hrs when levels stabilized. Serum radiolabel at
30 min appeared as conjugated and free (66:34). In the second study,
unlabelled E2 (1.5 mg) was injected (iv). Blood was taken at
0, 5, 15, 30, and 60 min and urine at 30 min. In serum, peak estrogen
levels appeared at 5 min (E2:E2C:E1C,
38:39:22 or 61:38 conjugated:free). Half-life of E2 is ca.10
min and 60 min for E2C. E1C peaked at 15 min and
declined by 20% in 45 min. Urine yielded similar results as the label
study (E2C:E1C, 60:40). Daily urine E2C
and weekly serum E2C and progesterone (P) were measured by
RIA in five mature female elephants. Preliminary results indicated
elevated levels of E2C during the luteal phase similar to
serum P profile. The follicular phase contains a biphasic profile of E2C.
These results suggest that E2 is rapidly converted to E2C
and excreted in urine.
Dahl, K.D.,
Czekala, N.M., Hsueh, A.J.W. Measurement of urinary bioactive
follicle-stimulating hormone (FSH) levels during reproductive cycles in
diverse mammalian species. Biology of Reproduction 40[suppl.1], 119.
1989.
Ref Type: Abstract
Abstract: Recently an in vitro granulosa cell aromatase bioassay (GAB)
was used to measure FSH in serum and urine samples (JCEM, March, 1987).
We now adapted the GAB assay to measure urinary bio-FSH levels in
conjunction with the determination of urinary immunoreactive
pregnanediol-3-glucoronide (PdG) and/or estrone conjugates (EC). Daily
urine samples were collected from 2 monkey species, lion-tailed macaque
(Macaca silenus) and golden monkey (Rhinopithecus roxellana), and 3
ungulates, giraffe (Giraffa camelopardalis), okapi (Okapi johnstoni) and
Asian elephant (Elephas maximus). The monkeys displayed a follicular
phase rise in EC followed by a midcycle bio-FSH surge which declined as
PdG concentrations increased during the luteal phase. Although both
monkey species had similar cycle lengths (30-35 days), the golden
monkey's follicular phase was much longer and displayed 2 bio-FSH peaks
compared to 1 peak for the macaque. Although EC was not detectable in
the ungulates, a midcycle FSH surge was followed by a luteal phase
increase in PdG. The closely related giraffe and okapi had similar
cycle lengths (16-17 days), follicular phase lengths (8-9 days), and
only 1 follicular phase FSH peak. Conversely, the elephants had cycles
lasting 16-19 weeks, and multiple FSH peaks were observed during the 5-6
week follicular phase. Thus, 1) in contrast to the single follicular
phase increase and preovulatory surge of bio-FSH seen in the macaque,
giraffe and okapi, the finding of multiple follicular phase peaks in the
golden monkey and elephants suggests the involvement of complex
regulatory mechanisms; 2) the GAB assay provides a noninvasive and
practical method for monitoring reproductive cycles in endangered
species for future breeding programs.
Morris, P.J.,
Held, J.P., Jensen, J.M. Clinical pathologic features of chronic renal
failure in an African elephant (Loxodonta africana).
Proc.1st.Intl.Conf.Zool.Avian Med. 468-472. 1987.
Ref Type: Conference Proceeding
Taylor-Robinson, D., Furr, P.M., Tully, J.G., Barile, M.F., Moller, B.R.,
1987. Animal models of Mycoplasma genitalium urogenital
infections. Israel Journal of Medical Sciences 23, 561-564.
Abstract: Male and female animals were inoculated urogenitally with
Mycoplasma genitalium, recovered originally from men with
nongonococcal urethritis. Mice, hamsters and male rhesus monkeys (Macaca
mulatta) were resistant. Male cynomolgus monkeys (Macaca
fascicularis) were not as sensitive as male chimpanzees (Pan
troglodytes): 9 of 11 developed an obvious genital tract infection,
some shedding organisms for more than 18 weeks. M. genitalium
was recovered from the blood of two of them when large numbers of
organisms were in the urethra. Most of the chimpanzees colonized with
the organisms had increased numbers of polymorphonuclear leukocytes in
the genital tract and developed a fourfold or greater antibody response.
Female squirrel monkeys (Saimiri sciureus) and female tamarins (Saguinus
mystar) exhibited low-level genital tract infections following
intravaginal inoculation, whereas marmosets (Callithrix jacchus)
and chimpanzees developed prolonged infections after similar
inoculation: thus, female chimpanzees shed organisms for 12 to 15
weeks. Marmosets and grivet monkeys (Ceropithecus aethiops)
developed salpingitis with antibody responses after intraoviduct
inoculation, and baboons (Papio anubis) developed parametritis
after intracervical inoculation. The results offer substantial evidence
for the pathogenicity of M. genitalium for the urogenital tract
of subhuman primates, and suggest that the microorganism may have a role
in human genital tract infections.
Jensen, J.
Paralumbar kidney biopsy in a juvenile African elephant.
Proc.Am.Assoc.Zoo Vet. 17. 1986.
Ref Type: Conference Proceeding
Abstract: Same case as Ref # 305.
Kahl, V.A.L.
The study of the reproductive cycle of the African elephant (Loxodonta
africana) through correlation of behavior and cytology with serum
and urinary hormonal patterns. 1986. Nebraska, Omaha, USA, University
of Nebraska.
Ref Type: Thesis/Dissertation
Munson, L.,
Heuschele, W., O'Banion, M.K., Sundberg, J.P., Oosterhuis, J.E., 1986.
Polyp in the urogenital canal of an African elephant. Journal of the
American Veterinary Medical Association 189, 1190-1191.
Poole, J.H.,
Kasman, L.H., Ramsay, E.C., Lasley, B.L., 1984. Musth and urinary
testosterone concentrations in the African elephant (Loxodonta
africana). Journal of Reproduction and Fertility 70, 255-260.
Abstract: Urine samples were obtained from free-ranging African
elephants that were considered to be in and out of musth. Testosterone
concentrations, measured by radioimmunoassay were significantly greater
in males that were in or around the time of behavioral musth. This
study supports a correlation between the observed behavioral
characteristics of musth and urinary testosterone levels.
Rasmussen,
L.E., Schmidt, M.J., Henneous, R., Groves, D., Daves, G.D.Jr., 1982.
Asian bull elephants: flehmen-like responses to extractable components
in female elephant estrous urine. Science 217, 159-162.
Abstract: Flehmen-like responses (urine tests) are one of the
characteristic behavioral reactions of male Asian elephants (Elephas
maximus) to cow elephants in estrus. Components of the urine of estrous
cow elephants were extracted with organic solvents and partially
purified by chromatography and shown to evoke Flehmen-like responses
when they were presented to adult bulls.
Ramsay, E.C.,
Lasley, B.L., Stabenfeldt, G.H., 1981. Monitoring the estrous cycle of
the Asian elephant (Elephas maximus), using urinary estrogens.
American Journal of Veterinary Research 42, 256-260.
Abstract: The estrous cycle of the Asian elephant (Elephas maximus)
was monitored by analysis of urinary estrogens. Daily morning urine
samples were analyzed for estrone (E1), estradiol (E2),
and total immunoreactive estrogen (ET). The ET
values were shown to correlate poorly with E1 and E2
and failed to reveal any patterns of reproductive cycling. Daily E1
and E2 values, indexed by creatinine concentrations,
demonstrated cyclic profiles in those samples of sufficient
concentrations. The technique offered a simple, noninvasive method for
determining ovarian function in the elephant.
Nirmalan, G.
Physiological peculiarties of elephants. State Level Workshop on
Elephants. 21-24. 1979. India, College of Veterinary and Animal
Sicences, Kerala Agricultural University.
Ref Type: Conference Proceeding
Fujimoto, E.,
Koto, N., Imori, T., Nakama, S., 1971. Gonadotropin in the urine of a
pregnant Indian elephant -- a case report. Zoologica 55, 73.
Abstract: In 1963, at Takarazuka Zoo, Japan, a young female Indian
elephant became pregnant, and in May, 1965, she gave birth to a very
large stillborn calf (weighing 133.3 kg, male). The time of conception
was problematical, but it was assumed as April or May of 1963, hence the
gestation period may have been 24 or 25 months, a little longer than
average. Pregnancy diagnosis was attempted during the early and middle
gestation period. For exploration, an urinary gonadotropin was checked
by the Friedman and Aschheim-Zondek tests on the whole urine samples
collected twice in August 1963. Results showed apparently positive
results in both tests. However, the samples collected in May and
September, 1964, showed negative in three tests, including a male frog (Rana)
reaction which was subjected to the concentrated urine samples. So,
probably a gondotropic substance many have been excreted in urine of
this elephant at some time of the early pregnancy, and this may be more
like FSH than LH in its activity.
McCullagh,
K.G., 1969. The growth and nutrition of the African elephant I. Seasonal
variations in the rate of growth and the urinary excretion of
hydroxyproline. East African Wildlife Journal 7, 85-90.
Orbanyi, J.,
Stohl, G., 1962. The urine of the elephant. UNKNOWN 321-328.
Evans, G.H.,
1961. Elephants and Their Diseases: A Treatise on Elephants. Government
Printing, Rangoon, Burma.
Simon, K.J.,
1959. Further studies on the urine of elephants. Indian Veterinary
Journal 36, 209-212.
Simon, K.J.,
1958. A preliminary study of the urine of elephants. Indian Veterinary
Journal 35, 345.
Schulte, T.L.,
1937. The genito-urinary system of the Elephas indicus male.
American Journal of Anatomy 61 , 131-157.
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.
Evans, G.H.,
1910. Elephants and Their Diseases: A Treatise on Elephants. Government
Printing, Rangoon, Burma.
Paterson,
A.M., 1898. The genito-urinary organs of the female Indian elephant.
Journal of Anatomy and Physiology 12, 582-604.
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.
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.
|