Seasonality of reproduction in sheep and its control by photoperiod

Seasonality of the reproductive cycle in sheep is a general phenomenon for mid-latitude breeds. The proximal part (breeding season) and also partially distal part (end of gestation and beginning of lactation) of this cycle is controlled by photoperiod, whatever the form of light regimens. Data are presented which indicate that male and female do not necessarily have the same photoperiodic sensitivity. Gonadal stimulation in the ram starts 1.5-2 months earlier than in the ewe under annual variations. Photoperiod controls the reproductive cycle by the intermediary of the hypothalamo-pituitary axis. There are both a steroid-independent and a steroid-dependent effect of light, depending on both decreasing and increasing daylength in mid-latitudes. Data are also presented which support Bunning's hypothesis on photoperiodic time measurement in mammals. Sheep measure photoperiodic time by using a circadian rhythm of photosensitivity. Daylength is not measured by the total duration of exposure to light but by the illumination of two special set points during the day, one of them entraining the circadian rhythm of photosensitivity and the other inducing or not inducing a physiological response if it is coincident, or not coincident, with photoinducible phase of that rhythm. A photoinducible phase has been found for prolactin secretion, and perhaps also for LH secretion. Melatonin secretion is used by sheep for measuring daylength. However, that secretion disappears during two set points during the day, thus raising the possibility of using alternatively melatonin and light pulse for controlling the reproductive cycle in sheep.     

Evidence for a circadian oscillation in the gonadal response of the tropical weaver bird (Ploceus philippinus) to programmed photoperiod.

The effect of an asymmetrical skeleton photoperiod scheduleee was studied on the gonadal development in a tropical finch, the weaver bird (Ploceus philippinus). The schedul comprised a short nonstimulatory primary photoperiod of 6 hrs and a secondary much shorter lightperiod given as a 15-min light pulse at different times in the dark period. The light pulse 11 hrs after the basic period resulted in gonadal stimulation, while light pulse in contimuation with the basic period or 8, 14 or 21 hrs after the basicperiod was not stimulatory. The "photoinducible phase" was much more precisely outlined by shifting the birds from an 8-h to a 10-h pulse, and from an 11-h to 12-h pulse and was found to be very short, lasting about an hour falling between 11 and 12 hrs after the primary light period. The short photoinducible phase may be of an adaptive value since in the tropics the difference between the shortest and the longest daylength is also rather small (3 hrs and 15 min at Varanasi, 25degrees N). Clearly the weaver bird possesses a fine time-measuring device involving an endogenous circadian rhythm in photosensitivity. In nature, spermatogenesis in this bird also begins in March when the daylength exceeds 11 hrs (thus perhaps coinciding with the photoinducible phase). In rather small, photoperiod may not serve as a cue to trigger seasonal reproductive periodicity, it seems that photoperiod can act as a Zeitgeber for the initiation of spermatogenesis in the weaver bird at least.     

Distribution and Prevalence of Parasitic Nematodes of Cowpea (Vigna unguiculata) in Burkina Faso

A comprehensive survey of the plant parasitic nematodes associated with cowpea (Vigna unguiculata) production fields was carried out in the three primary agro-climatic zones of Burkina Faso in West Africa. Across the three zones, a total of 109 samples were collected from the farms of 32 villages to provide a representative coverage of the cowpea production areas. Samples of rhizosphere soil and samples of roots from actively growing cowpea plants were collected during mid- to late-season. Twelve plant-parasitic nematode genera were identified, of which six appeared to have significant parasitic potential on cowpea based on their frequency and abundance. These included Helicotylenchus, Meloidogyne, Pratylenchus, Scutellonema, Telotylenchus, and Tylenchorhynchus. Criconemella and Rotylenchulus also had significant levels of abundance and frequency, respectively. Of the primary genera, Meloidogyne, Pratylenchus, and Scutellonema contained species which are known or suspected to cause losses of cowpea yield in other parts of the world. According to the prevalence and distribution of these genera in Burkina Faso, their potential for damage to cowpea increased from the dry Sahelian semi-desert zone in the north (annual rainfall < 600 mm/year), through the north-central Soudanian zone (annual rainfall of 600-800 mm/year), to the wet Soudanian zone (annual rainfall ≥ 1000 mm) in the more humid south-western region of the country. This distribution trend was particularly apparent for the endoparasitic nematode Meloidogyne and the migratory endoparasite Pratylenchus.     

Phylogeny and historical demography of economically important rodents of the genus Arvicanthis (Mammalia: Muridae) from the Nile Valley: of mice and men

The present study clarified the taxonomy, phylogeny and historical demography of semicommensal Nile rats ( Arvicanthis ) from the Nile Valley in Sudan. Nile rats are important crop pests and zoonotic disease reservoirs and are closely associated with agricultural settlements in the Nile Valley. Phylogenetic analysis of the complete cytochrome b gene (1140 bp) of 23 individuals from six localities in Sudan (from two previously recognized species, Arvicanthis niloticus and Arvicanthis testicularis ), supported the existence of only a single species, A. niloticus , from the Nile Valley. Historical demography of the Sudanese Nile Valley population inferred from mismatch coefficients indicated that an exponential population expansion event occurred approximately 144 000–288 000 years ago, corresponding in time with early human expansion and colonization from Africa to the Middle East, Europe, and the world. The inferred high level of gene flow and large size of Sudanese Nile populations of A. niloticus is consistent with historically recent (300 years ago) exponential human population growth and intense agricultural activity inferred from archaeological and historical evidence. Two African Arvicanthis clades were well supported by the broader phylogenetic analysis: (1) A. niloticus , Arvicanthis abyssinicus and Arvicanthis neumanni and (2) Arvicanthis rufinus and Arvicanthis ansorgei from western Africa. Within the first clade, divergence between lineages of A. niloticus s.s. from west and north-east Africa (8.9%) suggests specific recognition, but sampling of geographically intermediate localities is required. Based on hypothesized palaeodrainage and palaeoclimatic patterns, we propose a simple model for speciation of Arvicanthis in Africa.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 641–655.     

Parasitism of cowpea by Striga gesnerioides: variation in virulence and discovery of a new source of host resistance

In glasshouse experiments in the United Kingdom, Striga gesnerioides-resistant cowpea varieties, Suvita-2 and 58–57, were assessed alongside a susceptible variety for their responses to seventeen samples of S. gesnerioides from four West African countries. The susceptible variety was attacked by all samples, while Suvita-2 was resistant to samples from Burkina Faso and Mali, and 58–57 was resistant only to samples from Burkina Faso.
At least three physiological groups of the parasite can be distinguished: samples from Burkina Faso attacking the smallest range of cowpea varieties; those from Mali being intermediate; those from Niger and Nigeria attacking all the cowpea varieties. A sample from Cameroon behaved slightly differently and may represent a fourth type.
A cowpea line from Botswana, B.301, selected for its field resistance to Alectra vogelii proved resistant to all Striga populations to which it was exposed, including those from Niger and Cameroon.     

Tree mortality in the African Sahel indicates an anthropogenic ecosystem displaced by climate change

Aim Widespread reports of disappearing tree species and senescing savanna parklands in the Sahel have generated a vigorous debate over whether climate change or severe human and livestock pressure is principally responsible. Many of the tree taxa in decline are closely associated with human settlement and farming, suggesting that the parkland ecosystem may not be a natural vegetation assemblage. The aim of this study is to assess the possibility that human activities promoted the spread of taxa with edible fruit into dry Sudano‐Sahelian areas during high‐rainfall periods in the climate cycle. Location West African savannas (Mali, Burkina Faso, Ghana, Togo, Benin). Methods Cultivated savanna parklands and adjacent forests and transitional landscapes were inventoried at 27 sites in five countries. All trees with basal diameters > 10 cm were counted within 500‐m² belt transects. Species composition and abundance were contrasted between three landscape classes to assess the degree of influence exerted by traditional human management. Twentieth century rainfall data were averaged for two sets of weather stations encompassing the north–south range of typical parkland tree species. Rainfall trends were used to evaluate the putative impact of climate change on edible and/or succulent fruit species at the northern limit of the parkland savanna zone. Results Species composition and spatial distribution data indicate that the parkland ecosystem is significantly shaped by human activities. Indigenous land management favours edible‐fruit‐yielding taxa from the wetter Sudanian and Guinean vegetation zones over Sahelian species. Rainfall isohyets at the northern range limits of parkland species shifted southwards in the late 20th century, crossing the critical 600‐mm mean annual rainfall threshold for Sudanian flora. Relict vegetation and historical records indicate that the Sudanian parkland system extended in the past to near 15° N latitude in middle West Africa, compared with 13.5° N today. Main conclusions The current loss of mesic trees in the Sudano‐Sahel zone appears to be driven by the sharp drop in rainfall since the 1960s, which has effectively stranded anthropogenically distributed species beyond their rainfall tolerance limits.     

Effects of climate and local aridity on the latitudinal and habitat distribution of Arvicanthis niloticus and Arvicanthis ansorgei (Rodentia, Murinae) in Mali

Introduction The genus Arvicanthis (Lesson 1842) (Rodentia: Murinae), usually referred to as the unstriped grass rat, is mainly distributed in savanna and grassland habitats of Sub-Saharan Africa. Among the four chromosomal forms of Arvicanthis recently differentiated in Western and Central Africa, the one with a diploid chromosomal number (2n) of 62 and an autosomal fundamental number (NFa) of 62 or 64 is ascribed to Arvicanthis niloticus (Demarest 1822), while the one with 2n = 62 and a NFa between 74 and 76 is referred to A. ansorgei (Thomas 1910). Despite the broad area of sympatry recently uncovered along the inner delta of the Niger river in Mali [details in Volobouev et al. (2002) Cytogenetics and Genome Research, 96, 250–260], the distribution of the two species is largely parapatric and follows the latitudinal patterns of the West-African biogeographical domains, which are related to the latitudinal patterns of annual rainfall in this region. Here, we analyse the suggestion that the two species show specific adaptations to differences in climate aridity. Methods Karyologically screened animals were sampled in 19 localities in seasonally flooded regions located along the ‘Niger’ river in Mali and extending from 1100 to 200 mm of mean annual rainfall. The analysis of trapping success (TS) data allowed us to investigate the respective effects of climate (i.e. annual rainfall) and local (i.e. duration of the green herbaceous vegetation) aridity on the latitudinal and habitat distribution of the two species. Conclusions The broad zone of sympatry was found to correspond to a northward expansion of the recognized distribution area of A. ansorgei. TS values indicated that the two species responded very differently to climatic and local conditions of aridity. Arvicanthis ansorgei decreased in TS as regional conditions became more arid; a similar trend was also observed within regions where habitat occupancy decreased with local aridity. The higher TS observed in the most humid habitat relative to the others persisted throughout the latitudinal rainfall gradient. In contrast, TS of A. niloticus increased with latitudinal aridity. This species was present in more arid habitats than A. ansorgei from 1000 mm down to 400 mm of mean annual rainfall where a shift to the most humid habitat occurred. These opposite trends in TS distribution between species suggest that A. ansorgei is less adapted than A. niloticus to arid environments at both a regional and habitat level; thus, A. ansorgei would be able to invade dry regions only along the extensive floodplains bordering the inner delta of the ‘Niger’ river. Several biological traits that may be involved in limiting the southward distribution of A. niloticus are discussed.     

A circadian clock measures photoperiodic time in the male lizard Anolis carolinensis

Summary Photoperiod plays an important role in controlling the annual reproductive cycle of the male lizard Anolis carolinensis. The nature of photoperiodic time measurement in Anolis was investigated by exposing anoles to 3 different kinds of lighting paradigms (resonance, T cycles, and night breaks) to determine if photoperiodic time measurement involves the circadian system. Both the reproductive response and the patterns of entrainment of the activity rhythm were assessed. The results show that the circadian system is involved in photoperiodic time measurement in this species and that a discrete photoinducible phase resides in the latter half of the animals' subjective night. Significantly, the ability of the circadian system to execute photoperiodic time measurement is crucially dependent on the length of the photoperiod. Resonance, T cycle and night break cycles utilizing a photoperiod 10–11 h in duration reveal circadian involvement whereas these same cycles utilizing 6 or 8 h photoperiods do not.Abbreviation CRPP circadian rhythm of photoperiodic sensitivity     

Life-span growth dynamics and xylem anatomical patterns of diffuse-porous Afzelia africana Sm. (Fabaceae) in different ecological zones in Burkina Faso

Tropical West African savannas are exposed to high climatic variability with potential impacts on tree growth, forest dynamics and ecosystem productivity. In such context, understanding the long-term ecological responses of savanna trees to changing environmental conditions is of great relevance for taking appropriate conservation actions. We conducted the first study on tree-ring analysis and quantitative wood anatomy on Afzelia africana Sm. in Burkina Faso, to investigate the life-span growth trajectories and wood anatomical adjustment to site and to climate variations. A total of 24 stem discs was collected in four protected forests along the Sudano-sahelian and the Sudanian climatic zones. Wood samples were analyzed using standard dendrochronological methods and quantitative wood anatomy. The mean annual growth rates varied from 1.002 (± 0.249) mm. year⁻¹ in the Sudanian zone to 1.128 (± 0.436) mm. year⁻¹ in the Sudano-sahelian zone. Analysis of growth trajectories showed high variations within sites and between climatic zones. Wood anatomical traits significantly varied between sites. Principal Component Analysis revealed strong relationships between ring width, wood density and vessel traits, with 82.81 % of the total variance explained. Vessel size significantly increased from the pith to the bark, highlighting the ontogenetic effects on xylem anatomical variations. Inverse relationships were found between vessel size and vessel density across the driest site and the wettest site, suggesting that the higher the rainfall, the taller the tree, the larger vessel size, but the lower vessel density. By contrast, more arid conditions and high evapotranspiration lead to smaller vessel sizes and higher vessel density. Such anatomical adjustments highlight the trade-offs between water conductance efficiency and hydraulic safety, and emphasize physiological responses to climate variability. These variations on the long-term dynamics and xylem anatomical patterns underline complex interactions between ontogenetic effects and contrasting environmental factors that affect the eco-physiological functioning of A. africana throughout the Sudanian region.     

Regulation by Photoperiod of Seasonal Changes in Body Mass and Reproductive Function in Gray Mouse Lemurs (Microcebus murinus): Differential Responses by Sex

Microcebus murinus exhibits highly seasonal biological rhythms to cope with extreme seasonality in availability of resources. To study the role of daylength on seasonal changes in body mass and reproductive function, we exposed male and female gray mouse lemurs to natural, constant, or alternating light cycles for 2 years under constant environmental conditions. When exposed to either constant short (SD: 10 h light/day), long (LD: 14 h light/day), or intermediate (ID: 12 h light/day) daylength, males and females maintained a constant body mass with no spontaneous cyclic variation. We only observed typical seasonal body mass changes in subjects exposed to alternating periods of SD and LD, the weight gain being triggered by SD, whereas weight loss occurred under LD. Reproductive activity in females proceeded from an endogenous rhythm that was expressed under constant daylengths. In contrast, changes in reproductive activity in males depended on daylength variation. In both sexes, SD and LD have direct inhibitory or stimulatory effects on reproductive activity. In females, daylength regulates breeding season by synchronizing an endogenous sexual rhythm with the season, whereas in males, the perception of a critical photoperiod is used to determine the subsequent onset or arrest of their breeding season. These sexual differences in the effect of daylength could be related to sex-specific differences in reproductive constraints.     

Impact of environmental conditions on fruit production patterns of shea tree (Vitellaria paradoxa C.F.Gaertn) in West Africa

A better knowledge of the impact of environmental conditions on shea tree’s fruit production is critical to improve the species management. Fruit production of 237 shea trees was monitored over five successive years (2014–2018) to understand how climatic variables, shea tree population and land use could affect fruit production patterns of the species in Burkina Faso. Data were collected in 12 shea tree populations distributed in three climatic zones with two contrast land use systems. The results showed that fruit production was negatively correlated with mean annual temperature, temperature seasonality, rainfall seasonality and solar radiation, while it was positive with annual rainfall. Therefore, fruiting modelling is needed to predict the impact of future climate change on reproductive ability of the species. In each climatic zone, fruit production was significantly different among shea tree populations (p < 0.05). Within populations, fruit production greatly varies among years with specific patterns of variation. Land use had significant effect on fruit production of shea tree (p < 0.05). Annually, shea trees produced about 12 times more in agroforestry parklands than in protected areas. Interannual variation in fruit production at individual tree level was higher in protected areas. The cultivation of shea tree is recommended to improve its fruit productivity.     

The Drosophila circadian network is a seasonal timer

Previous work in Drosophila has defined two populations of circadian brain neurons, morning cells (M-cells) and evening cells (E-cells), both of which keep circadian time and regulate morning and evening activity, respectively. It has long been speculated that a multiple oscillator circadian network in animals underlies the behavioral and physiological pattern variability caused by seasonal fluctuations of photoperiod. We have manipulated separately the circadian photoentrainment pathway within E- and M-cells and show that E-cells process light information and function as master clocks in the presence of light. M-cells in contrast need darkness to cycle autonomously and dominate the network. The results indicate that the network switches control between these two centers as a function of photoperiod. Together with the different entraining properties of the two clock centers, the results suggest that the functional organization of the network underlies the behavioral adjustment to variations in daylength and season.     

Reproductive phase dependent circadian variations of plasma melatonin, testosterone, thyroxine and corticosterone in Indian palm squirrel, Funambulus pennanti

To explain the complex mechanism of environmental influence along with internal hormonal (factors) milieu on daily variations in the circulating levels of melatonin, testosterone, thyroxine and corticosterone were analyzed with the help of inferential statistics (Cosinor rhythmometry) in a seasonally breeding tropical rodent, F. pennanti during the reproductively active (RAP) and inactive phases (RIP). Plasma melatonin, thyroxine and corticosterone levels exhibited a significant circadian oscillation during both the active and inactive phases of the annual reproductive cycle. Melatonin showed higher amplitude during RIP in the circulating plasma. Testosterone presented a peak level during evening hours (16:00 - 18:00 h) during RAP only. The phase of thyroxine was noted ∼09:76 h and ∼10.35 h during active and inactive phases, respectively. Corticosterone showed a peak level at ∼12.00 h during both phases of the reproductive cycle. Further, in this tropical rodent, the minimum difference in photoperiod (∼3 - 4 hours) and maximum variation in temperature (max. 18°C - min. 10°C during RIP and max. 45°C - min. 32°C during RAP) and humidity (85% during RIP and 35% during RAP) regulated the diurnal rhythm of circulating melatonin circadian rhythm by ∼1 hour phase advance during RIP. In conclusion, the studied hormonal rhythms may be part of an integrative system to coordinate reproduction and physiological processes successfully with environmental factors.     

Night length measurements by the circadian clock controlling diapause induction in the mosquito Aedes atropalpus

Night interruption experiments were used to investigate the behavior of the clock controlling diapause induction in the mosquito, Aedes atropalpus. The data from these experiments indicated that the clock included a circadian oscillator which was phase set at dusk. Following this event the oscillator proceeded to drive a nightly rhythm of sensitivity to light. This rhythm included a photoinducible phase where light interruption inhibited diapause. The photoinducible phase was fixed, occurring 7 to 9 hr after dusk in all photoperiod regimens tested. The photoinducible phase was followed by a refractory phase, which continued until dawn. During the refractory period light did not inhibit diapause. These observations indicated that the circadian clock behaved like an interval timer which was set at dusk. The rhythm of sensitivity to light, an inherited time scale, limited the induction of diapause to seasonal periods when nights were longer than 9 hr. As a result, diapause was induced only when the daylength dropped below the critical photoperiod of L15:D9 (hours of light:hours of dark).A ‘T’ experimental design was used to confirm the importance of the length of the night in clock controlled induction of diapause in this mosquito.     

Circadian rhythm of sex pheromone-releasing behaviour in females of the dermestid beetle,Trogoderma glabrum: Regulation by photoperiod

The effects on sex pheromone-releasing, or calling behaviour, of diel photoperiods of varying daylength, of light cycle phase shifts, and of continuous illumination were investigated in Trogoderma glabrum females. On light régimes with 8 to 20 hr daylengths, calling maxima tended to centre close to photophase midpoints. Although influencing the time of day at which calling occurred, daylength had little effect on the amount of activity or the length of the calling period. When 16 : 8 LD light cycles were advanced or delayed by 4 hr, the time of day at which calling peaks were observed shifted within 2 to 4 cycles so that a constant phase relationship with photoperiod was maintained. Daily calling peaks were evident in groups of females exposed to between 1 and 5 days of continuous illumination, but mean calling time occurred earlier in the day as light exposures were lengthened. It was concluded that a circadian rhythm of calling behaviour exists in T. glabrum females. and that the rhythm can be entrained to 24 hr periodicity by photoperiod.     

Importance of photoperiodic signal quality to entrainment of the circannual reproductive rhythm of the ewe

An endogenous circannual rhythm drives the seasonal reproductive cycle of a broad spectrum of species. This rhythm is synchronized to the seasons (i.e., entrained) by photoperiod, which acts by regulating the circadian pattern of melatonin secretion from the pineal gland. Prior work has revealed that melatonin patterns secreted in spring/summer entrain the circannual rhythm of reproductive neuroendocrine activity in sheep, whereas secretions in winter do not. The goal of this study was to determine if inability of the winter-melatonin pattern to entrain the rhythm is due to the specific melatonin pattern secreted in winter or to the stage of the circannual rhythm at that time of year. Either a summer- or a winter-melatonin pattern was infused for 70 days into pinealectomized ewes, centered around the summer solstice, when an effective stimulus readily entrains the rhythm. The ewes were ovariectomized and treated with constant-release estradiol implants, and circannual cycles of reproductive neuroendocrine activity were monitored by serum LH concentrations. Only the summer-melatonin pattern entrained the circannual reproductive rhythm. The inability of the winter pattern to do so indicates that the mere presence of a circadian melatonin pattern, in itself, is insufficient for entrainment. Rather, the characteristics of the melatonin pattern, in particular a pattern that mimics the photoperiodic signals of summer, determines entrainment of the circannual rhythm of reproductive neuroendocrine activity in the ewe.     

Effect of thiopental sodium and barbitone sodium on the total acetylcholine content and acetylcholinesterase activity in the brain tissue of Arvicanthis niloticus

The total ACh content and AChE activity were determined 1 hr after the i.p. injection of different doses of thiopental sodium (5, 10 and 20 mg/ml/100 g body wt) and barbitone sodium (20, 40 and 80 mg/ml/100 g body wt). The effect of different time intervals (1 min, 10 min, 30 min, 1 hr, 2.5 hr, 5 hr, 8 hr, 12 hr, 24 hr and 48 hr) on the total ACh content and AChE activity was investigated after i.p. injection of 10 mg thiopental sodium and 40 mg barbitone sodium/ml/100 g body wt. Both thiopental sodium and barbitone sodium increased the total ACh content in the brain tissue of Arvicanthis niloticus. Both drugs inhibited the brain AChE activity. It is thought that the increase in the total ACh content in the brain tissue of Arvicanthis niloticus may be due to a decrease in the release of ACh from the neuronal tissue and a decrease in AChE activity.     

Control of annual gonadal cycles in Indian songbirds

Abstract

Reproduction is a part of life cycle with great environmental dependence. In contrast to temperate avian species, which mostly breed during summer, the Indian songbirds have more flexible breeding programs and exhibit a spectrum of reproductive strategies with the breeding season scattered all over the year. Control of breeding cycles in the Indian songbirds, therefore, are broadly viewed in light of two strategies (i) birds showing strong photoperiodic component in regulation of reproductive and post-reproductive events (ii) birds that do not exhibit typical photoperiodic regulation indicating the involvement of an inherent rhythm of reproduction. Both circadian and circannual rhythms have been demonstrated to regulate annual gonadal cycles of Indian songbirds. While photoperiod continues to be a predominant proximate factor for timing of breeding in majority of Indian songbirds investigated so far, some studies reveal the role of non photoperiodic cues such as the food availability, temperature, rainfall, etc. in timing/modulating the timing of breeding. The conversion or non-conversion of thyroxine to triiodothyronine may act as a long or short photoperiod signal and may up or downregulate the synthesis and release of GnRH-I in hypothalamus, FSH and LH in anterior pituitary and gonadal steroids in gonads causing gonadal growth or regression, respectively.     

Latitudinal clines in the properties of a circadian pacemaker

The circadian rhythm of eclosion activity and its pacemaker were analyzed in a series of latitudinal races of Drosophila auraria ranging from 34.2 degrees to 42.9 degrees N in Japan. The phase of the rhythm (psi EL) to the daily photoperiod (PP) changes as daylength is increased, and the slope of psi EL (PP) changes with latitude. Is is sufficiently greater in the north to cause a phase reversal of northern and southern races on long versus short photoperiods. This reversal is found in assays of the pacemaker's phase (psi PL) as well as that of the rhythm (psi EL). Assay of the pacemaker shows that its period (tau) is longer in northern than in southern races, and that the amplitude of its phase response curve (PRC) is lower in the north. The period of the rhythm in all latitudinal races is longer than 24 hr in short photoperiods (LD 1:23), but is probably less than 24 hr (as an aftereffect of photoperiod) in longer days such as LD 14:10. The observed north-south differences in the phase relation of both pacemaker and rhythm to the light cycle are explained by the latitudinal clines in pacemaker properties and a postulated aftereffect of photoperiod on tau. It is suggested that the latitudinal cline in PRC amplitude has functional significance in conserving the amplitude of the pacemaker's signal to the rest of the system it times. Computer simulation shows that without such a reduction in the perceived light intensity, pacemaker amplitude will be lowered by the increase in duration of the daily light at higher latitudes.     

The circadian timing system and reproduction in mammals.

Circadian systems in a wide variety of organisms all appear to include three basic components: 1) biological oscillators that maintain a self-sustained circadian periodicity in the absence of environmental time cues; 2) input pathways that convey environmental information, especially light cues, that can entrain the circadian oscillations to local time; and 3) output pathways that drive overt circadian rhythms, such as the rhythms of locomotor activity and a variety of endocrine rhythms. In mammals, the circadian system is employed in the regulation of reproductive physiology and behavior in two very important ways. 1) In some species, there is a strong circadian component in the timing of ovulation and reproductive behavior, ensuring that these events will occur at a time when the animal is most likely to encounter a potential mate. 2) Many mammals exhibit seasonal reproductive rhythms that are largely under photoperiod regulation; in these species, the circadian system and the pineal gland are crucial components of the mechanism that is used to measure day length. The rhythm of pineal melatonin secretion is driven by a neural pathway that includes the circadian oscillator(s) in the suprachiasmatic nuclei. Melatonin is secreted at night in all mammals, and the duration of each nocturnal episode of melatonin secretion is inversely related to day length. The pineal melatonin rhythm appears to serve as an internal signal that represents day length and that is capable of regulating a variety of seasonal variations in physiology and behavior.