Seasonal cycles in the blood plasma concentration of FSH, inhibin and testosterone, and testicular size in rams of wild, feral and domesticated breeds of sheep

Seasonal cycles in testicular activity in rams were monitored in groups of wild (mouflon), feral (Soay) and domesticated breeds of sheep (Shetland, Blackface, Herdwick, Norfolk, Wiltshire, Portland and Merino) living outdoors near Edinburgh (56 degrees N). The changes in the blood plasma concentrations of FSH, inhibin and testosterone, and the diameter of the testis were measured every half calendar month from 1 to 3 years of age. There were significant differences between breeds in the magnitude and timing of the seasonal reproductive cycle. In the mouflon rams, the seasonal changes were very pronounced with a 6-15-fold increase in the plasma concentrations of FSH, inhibin and testosterone from summer to autumn, and a late peak in testicular diameter in October. In the Soay rams and most of the domesticated breeds, the seasonal increase in the reproductive hormones occurred 1-2 months earlier with the peak in testicular size in September or October. In the two southern breeds (Portland and Merino), the early onset of testicular activity was more extreme with the seasonal maximum in August. In cross-bred rams, produced by mating Soay ewes (highly seasonal breed) with Portland or Merino rams (less seasonal breeds), there was a seasonal reproductive cycle that was intermediate compared to that of the parents. A comparison between all 11 breeds showed a significant correlation between the timing of the seasonal cycle in plasma FSH concentration and testicular diameter (time of peak FSH vs testis, r = 0.95). The overall results in the rams are consistent with a primary role of FSH in dictating the seasonal cycle in testicular size and the secretion of inhibin. The earlier seasonal onset in the testicular cycle in the southern breeds of domesticated sheep, and the differences from the wild type, are taken to represent the effects of genetic selection for a longer mating season.     

Seasonal cycles in testicular activity in Mouflon, Soay sheep and domesticated breeds of sheep: breeding seasons modified by domestication

The seasonal reproductive cycle in rams was monitored in Mouflon (wild-type), Soay (feral type) and a variety of domesticated breeds of sheep (Shetland, Blackface, Herdwick, Norfolk, Wiltshire, Portland and Merino) by measuring the changes in the diameter of the testes (first three years of life in all breeds) and the blood plasma concentrations of FSH and testosterone (first four to six years of life in Mouflon and Soay rams). In the Mouflon rams there was a pronounced seasonal cycle in all the reproductive parameters from one to six years of age. The plasma concentrations of FSH increased from June to September associated with redevelopment of the testes; maximum testicular size and plasma concentrations of testosterone occurred in October at the onset of the rut. In the Soay and domesticated breeds the seasonal maximum in testicular size occurred in late September or October except in two of the southern breeds (Portland and Merino) which showed an earlier peak to the sexual cycle in August. The change in size of the testes from the minimum to the maximum was less in the domesticated breeds (135–157%) compared to the Soay (171%) and Mouflon (160%). Crossbred rams produced by mating Soay ewes (highly seasonal breed) with Portland or Merino rams (less seasonal breed) had a seasonal testicular cycle intermediate in timing compared to the pattern characteristics of the parent breeds; this is consistent with the involvement of multiple genes in the mechanism controlling the sexual cycle in the ram. The earlier seasonal onset of full testicular activity in the southern breeds is assumed to be the result of selection for a prolongation of the breeding season for early lambing.     

Influence of age on the relationship between annual changes in horn growth rate and prolactin secretion in the European mouflon (Ovis gmelini musimon)

Annual variations in the growth of horns, and their correlation with seasonal changes of testicular size, and prolactin (PRL) and melatonin secretion were monitored in six pubertal mouflon rams living in their original latitude (40 degrees N). Mouflons born and maintained under captive conditions were classified in two age classes: sub-adult (2 years; n=3) and adult (> or =3 years; n=3). The rate of horn growth was greater (P <0.001) in sub-adult than in adult mouflon rams. Horn growth was influenced by season in both adult and sub-adult mouflons (P <0.05) with largest monthly growth occurring in spring and summer. Seasonal variations of plasma PRL concentrations were correlated with horn growth in adult, but not in sub-adult mouflon rams. The rate of horn growth was inversely correlated with testicular size (r=-0.5, P=0.07). Seasonal changes in the amplitude of the daily melatonin rhythm in solstices and equinoxes were observed, which were not correlated with variations in the rate of horn growth. These results provide support for a possible role of PRL in the control of growth of horns in the adult mouflon.     

Seasonality of semen production and plasma luteinizing hormone,testosterone and prolactin levels in Romney,Merino and polled dorset rams

An attempt was made to define the seasonality of reproduction in rams in the southern hemisphere by repeated measurement of semen characteristics and of plasma luteinizing hormone (LH), testosterone and prolactin concentrations. These parameters were studied for 16 months in Romney rams on pasture, with Merino and Polled Dorset rams included for comparison.Semen from all three breeds showed regular seasonal changes in ejaculate volumes, with peak values being recorded during March. A similar autumnal peak of seminal fructose levels was noted for ejaculates from Romney and Merino rams, but not for those from Polled Dorsets. Most of the other semen parameters measured showed little tendency for seasonal variations. However, a change in semen collection technique, from predominantly artificial vagina to entirely electroejaculation, may have masked some seasonal changes.Plasma hormone levels also varied in a regular manner, with peak levels occurring in summer and autumn: highest levels for prolactin were recorded in November–March, for LH in December–February and for testosterone in January–March. An exception to this pattern was recorded from the Merino rams, for which there was no definite peak of LH secretion.It is suggested that these seasonal changes resulted primarily from changes in daily photoperiod.     

Nocturnal variation of prolactin secretion in the Mouflon (Ovis gmelini musimon) and domestic sheep (Ovis aries): seasonal changes

Seasonal changes in nocturnal prolactin secretion and their relationship with melatonin secretion were monitored in wild (Mouflon, Ovis gmelini musimon) and domesticated sheep (breed Manchega, Ovis aries). Two groups of eleven adult females each, were maintained outdoors under natural photoperiod. Plasma concentrations of prolactin and melatonin were determined during the summer and winter solstices and the autumn and spring equinoxes. Blood samples were collected every 3h during the night hours, and 1h before and after the onset of darkness and sunrise. Maximum mean plasma concentrations of prolactin during the dark-phase in Mouflons were observed in the summer solstice, (P<0.001) and in the summer solstice and spring equinox in Manchega ewes (P<0.001). Mean plasma concentrations of prolactin were higher in the wild species (P<0.001) during the summer solstice. In contrast, during the spring equinox, mean levels of prolactin were higher in Manchega ewes than in Mouflons (P<0.05). Plasma prolactin concentrations showed a nocturnal rhythm in both breeds, with seasonal variations (P<0.001). The increase in plasma melatonin levels during the first hour after sunset was accompanied to increasing concentrations of PRL 1h after the onset of darkness, only in the autumn and spring equinox for the Mouflon, and in the summer solstice and spring equinox for the Manchega ewes. In Mouflons, the fall of plasma PRL concentrations about the middle dark-phase in all the periods studied, coincided with high levels of melatonin. A similar relation was observed in Manchega ewes only in the winter solstice and spring equinox. The current study shows that the nocturnal rhythm of prolactin secretion exhibits seasonal variation; differences in the patterns of prolactin secretion between Mouflon and Manchega sheep are taken to represent the effects of genotype.     

Daylength influences pelage and plasma prolactin concentrations but not reproduction in the prairie vole, Microtus ochrogaster

Short daylengths did not affect testes weight or spermatogenic index in male voles or uterine weight in female voles. Short daylengths did stimulate the growth of a winter pelage in both sexes; short-day voles had longer underhairs and guard hairs and a thicker, more dense pelage than did long-day voles. Plasma prolactin concentrations were five times higher in long-day than in short-day females and 25% higher in long-day males than in short-day males. The effect of short daylength on pelage was prevented by pinealectomy. We suggest that the growth of a winter coat is an obligate adaptation for winter survival, stimulated by exposure to short daylengths, but that changes in breeding activity are facultative and dependent to a greater extent on other cues for seasonal synchronization.     

Introgression and the fate of domesticated genes in a wild mammal population

When domesticated species are not reproductively isolated from their wild relatives, the opportunity arises for artificially selected variants to be re‐introduced into the wild. However, the evolutionary consequences of introgression of domesticated genes back into the wild are poorly understood. By combining high‐throughput genotyping with 25 years of long‐term ecological field data, we describe the occurrence and consequences of admixture between a primitive sheep breed, the free‐living Soay sheep of St Kilda, and more modern breeds. Utilizing data from a 50 K ovine SNP chip, together with forward simulations of demographic scenarios, we show that admixture occurred between Soay sheep and a more modern breed, consistent with historical accounts, approximately 150 years ago. Haplotype‐sharing analyses with other breeds revealed that polymorphisms in coat colour and pattern in Soay sheep arose as a result of introgression of genetic variants favoured by artificial selection. Because the haplotypes carrying the causative mutations are known to be under natural selection in free‐living Soay sheep, the admixture event created an opportunity to observe the outcome of a ‘natural laboratory’ experiment where ancestral and domesticated genes competed with each other. The haplotype carrying the domesticated light coat colour allele was favoured by natural selection, while the haplotype associated with the domesticated self coat pattern allele was associated with decreased survival. Therefore, we demonstrate that introgression of domesticated alleles into wild populations can provide a novel source of variation capable of generating rapid evolutionary changes.     

Orchestration of avian reproductive effort: an integration of the ultimate and proximate bases for flexibility in clutch size, incubation behaviour, and yolk androgen deposition

How much effort to expend in any one bout of reproduction is among the most important decisions made by an individual that breeds more than once. According to life-history theory, reproduction is costly, and individuals that invest too much in a given reproductive bout pay with reduced reproductive output in the future. Likewise, investing too little does not maximize reproductive potential. Because reproductive effort relative to output can vary with predictable and unpredictable challenges and opportunities, no single level of reproductive effort maximizes fitness. This leads to the prediction that individuals possessing behavioural mechanisms to buffer challenges and take advantage of opportunities would incur fitness benefits. Here, we review evidence in birds, primarily of altricial species, for the presence of at least two such mechanisms and evidence for and against the seasonal coordination of these mechanisms through seasonal changes in plasma concentrations of the pituitary hormone prolactin. First, the seasonal decline in clutch size of most bird species may partially offset a predictable seasonal decline in the reproductive value of offspring. Second, establishing a developmental sibling-hierarchy among offspring may hedge against unpredictable changes in resource availability and offspring viability or quality, and minimize energy expenditure in raising a brood. The hierarchy may be a product, in part, of the timing of incubation onset relative to clutch completion and the rate of yolk androgen deposition during the laying cycle. Because clutch size should influence the effects of both these traits on the developmental hierarchy, we predicted and describe evidence in some species that females adjust the timing of incubation onset and rate of yolk androgen deposition to match clutch size. Studies on domesticated precocial species reveal an inhibitory effect of the pituitary hormone prolactin on egg laying, suggesting a possible hormonal basis for the regulation of clutch size. Studies on the American kestrel (Falco sparverius) and other species suggest that the seasonal increase in plasma concentrations of prolactin may regulate both a seasonal advance in the timing of incubation onset and a seasonal increase in the rate of yolk androgen deposition. These observations, together with strong conceptual arguments published previously, raise the possibility that a single hormone, prolactin, functions as the basis of a common mechanism for the seasonal adjustment of reproductive effort. However, a role for prolactin in regulating clutch size in any species is not firmly established, and evidence from some species indicates that clutch size may not be coupled to the timing of incubation onset and rate of yolk androgen deposition. A dissociation between the regulation of clutch size and the regulation of incubation onset and yolk androgen deposition may enable an independent response to the predictable and unpredictable challenges and opportunities faced during reproduction.     

Horn growth related to testosterone secretion in two wild Mediterranean ruminant species: the Spanish ibex (Capra pyrenaica hispanica) and European mouflon (Ovis orientalis musimon)

Seasonal variations in the horn development and testicular activity of the Spanish ibex (Capra pyrenaica hispanica) (n=6) and European mouflon (Ovis orientalis musimon) (n=5) were monitored to determine the role of increasing testosterone concentration on the arrest of horn growth during the rutting season. Marked seasonal variations in the rate of horn growth (P<0.01) and testicular activity (P<0.001) were seen in both species, although the magnitude and timing of these changes were different (P<0.01). Horn growth rate was inversely correlated to seasonal levels in testosterone plasma concentration in both species (ibex: R=-0.45, P<0.01; mouflon R=-0.51, P<0.01). In the mouflon, the increase in plasma testosterone concentration recorded in September (P<0.05 compared with the lowest concentration) coincided with a significant reduction in horn growth (P<0.05). In the ibex, the increase in plasma testosterone concentration in October (P<0.05 compared with the lowest concentration) was associated with a significant arrest of horn growth in November (P<0.05). These results appear to support the hypothesis that high peripheral plasma levels of testosterone are linked with the seasonal arrest of horn growth during the rutting period.     

Prolactin-dependent seasonal changes in pelage: role of the pineal gland and dopamine.

The Siberian hamster displays seasonal changes in pelage that are dependent upon fluctuations in circulating prolactin levels. Pinealectomy prevented the decrease in serum prolactin and molt to the winter pelage displayed by castrated males housed under a short-day photoperiod. A dopaminergic antagonist, pimozide, enhanced prolactin levels in both pinealectomized and sham-operated animals under both long and short photoperiods. In the short-day animals, this effect of pimozide was associated with a prevention of the development of winter pelage. These results indicate that seasonal prolactin levels and related pelage changes are dependent upon the integrity of the pineal gland. However, basal prolactin levels under different photoperiod conditions appear to be only partly regulated by the actions of the dopaminergic system.     

Ovulatory activity and plasma prolactin concentrations in wild and domestic ewes exposed to artificial photoperiods between the winter and summer solstices

Mouflon and domestic Manchega sheep differ in the timing of their reproductive season under natural photoperiod (NP) conditions. This study examines whether they also differ in their reproductive responses to artificial photoperiod cues. For this, mouflons (n=24) and ewes (n=24) were exposed between the winter and summer solstices to artificial long days (LD: 16 h light/day), to short days (SD) simulated via the use of melatonin implants, or to NP conditions (controls), and their ovulatory activity monitored. The effects of these treatments on annual changes in prolactin concentration were also recorded. In the LD mouflon ewes, the offset and onset (7 March ± 5 and 2 October ± 4, respectively) of cyclic ovulatory activity occurred earlier (P<0.001) than in the NP animals (26 April ± 6 and 20 October ± 2, respectively), but no differences were seen (P>0.05) between the SD and NP mouflon ewes in either the onset of anoestrus (12 May ± 6 and 26 April ± 6, respectively) or the onset of subsequent ovulatory activity (13 October ± 8 and 20 October ± 2, respectively); however the duration of the anoestrus period was significantly reduced in the SD. In LD Manchega ewes, the onset of anoestrus was advanced (2 February ± 5 vs 15 March ± 11), but ovulatory activity started at the same time as in NP Manchega ewes (16 July ± 4 vs 5 July ± 8). In the SD Manchega ewes, two animals showed continuous cyclic ovulatory activity over the course of the experiment while the remainder entered anoestrus two months later (16 May ± 6, P<0.001) than their NP counterparts. In these SD ewes, the onset of cyclic ovarian activity was very variable. An annual rhythm of plasma prolactin concentration was seen in both the mouflon and Manchega ewes under all three photoperiod conditions. However, the amplitudes of the changes seen in prolactin concentration were smaller in both the LD and SD animals than in the corresponding NP animals (P<0.001). In conclusion, the results show that these two types of Mediterranean sheep differ in their ovulatory response when subjected to artificial photoperiods. The results also suggest that refractoriness to SDs may be the most important physiological mechanism regulating the onset of anoestrus in highly seasonal breeds, but not in less seasonal breeds.     

Prolactin inhibition does not influence horn growth in two wild caprinae species: European mouflon (Ovis orientalis musimon) and Iberian ibex (Capra pyrenaica)

Studies on seasonal horn development and the endocrine mechanism regulating its pattern in wild ruminants are scarce. The aim of this paper was to study the influence of photoperiod and prolactin (PRL) on horn growth in two wild ruminant species: the European mouflon and the Iberian ibex. Eighteen male ibexes and 13 mouflon rams, maintained in captivity, were divided into three groups: a control group, kept under a natural photoperiod (latitude, 40°25′ N); a long-day group, exposed to an artificial photoperiod of 15-h light and 9-h darkness; and a group treated with bromocriptine (BCR; 10?mg twice weekly during spring and summer) to induce hypoprolactinaemia. Horn length growth (HLG) was recorded weekly for 18?months; plasma PRL concentrations were measured twice monthly by radioimmunoassay. In the ibexes of the long-day group, the period of strong horn development during spring–summer was significantly reduced by 2?months compared with the controls. In the mouflons of the long-day group, this same period was significantly increased by 9?months. In the BCR-treated animals, hypoprolactinaemia was observed in both species, but HLG was the same as in the corresponding controls. The present results suggest that the seasonal pattern of horn growth of wild ruminants is primarily modulated by photoperiod in a species-dependent manner. The persistence of resurgence of horn growth during spring in the BCR-induced hypoprolactinaemic animals of both species suggests that annual variations in blood PRL concentration have no effect on seasonal variation in horn growth.     

Seasonal changes in ovulatory activity, plasma prolactin, and melatonin concentrations, in mouflon (Ovis gmelini musimon) and Manchega (Ovis aries) ewes

Seasonal changes in ovulatory activity, plasma prolactin and melatonin concentrations were monitored in a wild (Mouflon) and a domesticated (Manchega) breed of sheep, both originating and living under similar latitudes (40 degrees N). Mouflons express ovarian cycles significantly later than Manchega ewes (October vs. July, P < 0.001); however, they ended cycling one month later than Manchegas (April vs. March, P < 0.05). While prolactin concentrations were high when Manchega ewes started to cycle, they were at their lowest concentrations when Mouflons started cycling. Overall, mean prolactin concentrations were higher (P < 0.001) in Mouflons than Manchegas throughout most of the year. Within the limits of sampling frequency, the duration of melatonin secretion was similar in both groups during the solstice and equinox periods; however, the amplitude was lower (P < 0.01) in Mouflons than Manchegas during the solstice periods. The significant breed differences in the seasonal hormonal changes may be attributed to a genetic influence in the endocrine responses to the same photoperiodic cues.     

Hormonal regulation of the annual pelage color cycle in the Djungarian hamster, Phodopus sungorus. II. Role of prolactin

This study investigated whether photoperiod-induced changes in circulating prolactin levels, which have been observed in the Djungarian hamster ( Yellon and Goldman, '83; Duncan and Goldman, ' 83a ), might be involved in seasonal pelage color changes in this species. Injection of ovine prolactin (100 micrograms/day) inhibited the short photoperiod-induced winter molt. This finding indicated that the suppression of endogenous prolactin levels normally occurring in short photoperiod-housed hamsters (Duncan and Goldman, ' 83a ) may induce the winter molt. Suppression of prolactin secretion with bromoergocryptine (200 micrograms/day) strongly inhibited the spring molt, while concomitant treatment with ovine prolactin (100 micrograms/day) overcame this effect of bromoergocryptine. Injection of bromoergocryptine (200 micrograms/day) stimulated the winter molt in castrated hamsters housed in long photoperiod; concomitant injection of prolactin (100 micrograms/day) reversed this effect as well. These findings strongly suggested that an increase in endogenous prolactin levels may be necessary for the development and maintenance of the summer pelage.     

Prolactin replacement fails to inhibit reactivation of gonadotropin secretion in rams treated with melatonin under long days

This study tested the hypothesis that prolactin (PRL) inhibits gonadotropin secretion in rams maintained under long days and that treatment with melatonin (s.c. continuous-release implant; MEL-IMP) reactivates the reproductive axis by suppressing PRL secretion. Adult Soay rams were maintained under long days (16L:8D) and received 1) no further treatment (control, C); 2) MEL-IMP for 16 wk and injections of saline/vehicle for the first 8 wk (M); 3) MEL-IMP for 16 wk and exogenous PRL (s.c. 5 mg ovine PRL 3x daily) for the first 8 wk (M+P). The treatment with melatonin induced a rapid increase in the blood concentrations of FSH and testosterone, rapid growth of the testes, an increase in the frequency of LH pulses, and a decrease in the LH response to N-methyl-D,L-aspartic acid. The concomitant treatment with exogenous PRL had no effect on these reproductive responses but caused a significant delay in the timing of the sexual skin color and growth of the winter pelage. These results do not support the hypothesis and suggest that PRL at physiological long-day concentrations, while being totally ineffective as an inhibitor of gonadotropin secretion, acts in the peripheral tissues and skin to maintain summer characteristics.     

Merino and Merino-derived sheep breeds: a genome-wide intercontinental study

Background

Merino and Merino-derived sheep breeds have been widely distributed across the world, both as purebred and admixed populations. They represent an economically and historically important genetic resource which over time has been used as the basis for the development of new breeds. In order to examine the genetic influence of Merino in the context of a global collection of domestic sheep breeds, we analyzed genotype data that were obtained with the OvineSNP50 BeadChip (Illumina) for 671 individuals from 37 populations, including a subset of breeds from the Sheep HapMap dataset.

Results

Based on a multi-dimensional scaling analysis, we highlighted four main clusters in this dataset, which corresponded to wild sheep, mouflon, primitive North European breeds and modern sheep (including Merino), respectively. The neighbor-network analysis further differentiated North-European and Mediterranean domestic breeds, with subclusters of Merino and Merino-derived breeds, other Spanish breeds and other Italian breeds. Model-based clustering, migration analysis and haplotype sharing indicated that genetic exchange occurred between archaic populations and also that a more recent Merino-mediated gene flow to several Merino-derived populations around the world took place. The close relationship between Spanish Merino and other Spanish breeds was consistent with an Iberian origin for the Merino breed, with possible earlier contributions from other Mediterranean stocks. The Merino populations from Australia, New Zealand and China were clearly separated from their European ancestors. We observed a genetic substructuring in the Spanish Merino population, which reflects recent herd management practices.

Conclusions

Our data suggest that intensive gene flow, founder effects and geographic isolation are the main factors that determined the genetic makeup of current Merino and Merino-derived breeds. To explain how the current Merino and Merino-derived breeds were obtained, we propose a scenario that includes several consecutive migrations of sheep populations that may serve as working hypotheses for subsequent studies.

Electronic supplementary material

The online version of this article (doi:10.1186/s12711-015-0139-z) contains supplementary material, which is available to authorized users.     

Coat colour inheritance in Soay, Orkney and Shetland sheep

Analysis of the numerical proportions of Soay, Orkney and Shetland sheep of different colours together with test matings, produced results compatible with the hypothesis that these breeds have a multiple allelic series at locus A , white ( A ₁) being dominant to grey ( A ₂) and both being dominant to the gene for self-colour ( A ₅). The alleles at the A locus are epistatic to the alleles for pigment production at locus B , black ( B ₁) being dominant to brown ( B ₂).     

Effects of breed, ovarian steroids and season on the pulsatile secretion of LH in ovariectomized ewes

The effects of season and of oestradiol and progesterone on the tonic secretion of LH were studied in ovariectomized Merino and Suffolk ewes, two breeds which differ markedly in the seasonal pattern of their reproductive activity. In the absence of exogenous steroids, the frequency of LH pulses was lower and the amplitude of the pulses was higher in anoestrus than in the breeding season for Merino and Suffolk ewes 30 days after ovariectomy. In long-term (190 days) ovariectomized ewes, this seasonal change in LH secretion was observed in Suffolk ewes only. During seasonal anoestrus, treatment of ewes with subcutaneous oestradiol-17 beta implants (3, 6 or 12 mm in length) decreased the frequency of LH pulses in a dose-dependent manner, with Suffolk ewes being far more sensitive to the inhibitory effects of oestradiol than Merino ewes. The lowest dose of oestradiol (3 mm) had no effect on the secretion of LH in Merino ewes, but reduced secretion in Suffolk ewes. Treatment of ewes with the highest dose of oestradiol (12 mm) completely abolished LH pulses in Suffolk ewes, whereas infrequent pulses remained evident in Merino ewes. During the breeding season, oestradiol alone had no effect on the pulsatile release of LH in either breed, but in combination with progesterone there was a significant reduction in LH pulse frequency. Progesterone effectively decreased LH secretion in both breeds in both seasons. It was concluded that differences between breeds in the 'depth' of anoestrus could be related to differences in the sensitivity of the hypothalamus to both negative feedback by oestradiol and the direct effects of photoperiod.     

Hormonal regulation of the annual pelage color cycle in the Djungarian hamster, Phodopus sungorus. I. Role of the gonads and pituitary

The Djungarian hamster exhibits an agouti pelage in the summer and a predominantly white pelage in the winter. This pelage color cycle is known to be regulated by the length of the daily photoperiod probably acting through the pineal gland, as is the seasonal cycle of reproductive function with which it is closely correlated ( Figala et al., '73; Hoffmann, ' 78b ). The possibility of a causal relationship between the decline in gonadal hormone secretion and the coat color change occurring in short photoperiod was examined. Gonadectomized and intact male and female hamsters were exposed to either long (16L:8D) or short ( 10L : 14D ) photoperiod for several months. Gonadectomy neither induced the change to the winter pelage color in long photoperiod-housed animals, nor prevented either the change to the winter pelage or the spontaneous return to summer pelage color in short photoperiod-housed animals. Chronic implants of testosterone in castrated males delayed and attenuated the short photoperiod-induced coat color change. Administration of ovine prolactin (100 micrograms/day) stimulated pigmentation in hamsters with the winter pelage, whereas administration of a alpha MSH (30 micrograms/day) was without effect. These results suggest that changes in pelage color may be regulated largely by changes in pituitary prolactin secretion and modified to some extent by changes in gonadal steroid hormone secretion.     

Seasonal changes in the reproductive organs and plasma and pituitary hormone content of the male Bennett's wallaby (Macropus rufogriseus vufogriseus)

In a number of species of seasonally breeding marsupial, the male is fertile throughout the year but there is a marked seasonal change in weight of the accessory sexual glands. In this study, body weight, prostate, epididymis and testis weights and plasma concentrations of testosterone, LH and prolactin and pituitary content of LH and prolactin were determined in male Bennett's wallabies shot at 1–2 month intervals over a period of 17 months. There was a highly significant increase in prostate weight which was coincident with the breeding season for this species. A small but significant increase in testis weight was also observed but epididymis weight remained unchanged. Plasma testosterone concentrations were significantly increased at a time coincident with the increase in prostate weight. Plasma prolactin and LH concentrations were low in most animals and remained unchanged during the study. In contrast, pituitary prolactin and LH contents showed highly significant changes, with increasing and peak hormone content preceding maximum prostate weight and plasma testosterone concentrations by several months. While these latter results suggest a role for prolactin and LH in the seasonal control of the reproductive organs in the male wallaby, a more intensive study of the pattern of secretion of these hormones and possibly more sensitive hormone assays are required to understand their relative roles in regulating the annual cycle of prostate growth.