Proximate developmental mediators of sexual dimorphism in size: case studies from squamate reptiles

Sexual dimorphism in size (sexual size dimorphism; SSD) is nearlyubiquitous, but the relative importance of genetic versus environmentalcontrol of SSD is not known for most species. We investigatedproximate determinants of SSD in several species of squamatereptiles, including three species of Sceloporus lizards andthe diamond-backed rattlesnake (Crotalus atrox). In naturalpopulations of these species, SSD is caused by sexual differencesin age-specific growth. Males and females, however, may oftenshare similar potentials for growth: growth is strongly responsiveto the availability of food, and sexual differences in growthcan be greatly suppressed or completely absent under commonenvironmental conditions in the laboratory. Sexually divergentgrowth is expressed in natural environments because of inherentecological differences between males and females and becauseof potential epigenetic effects of sex-specific growth regulators.In field-active Sceloporus, sexual differences in growth rateare associated with sexual divergence in plasma testosterone.Experiments confirm that testosterone inhibits growth in speciesin which females are larger (for example, S. undulatus and S.virgatus) and stimulates growth in those in which males arelarger (for example, S. jarrovii). Interestingly, however, sexualdivergence in plasma testosterone is not accompanied by divergencein growth in S. jarrovii or in male-larger C. atrox in the laboratory.Furthermore, experimental effects of castration and testosteronereplacement on growth are not evident in captive S. jarrovii,possibly because growth effects of testosterone are supersededby an abundant, high-quality diet. In female-larger S. undulatus,growth may be traded-off against testosterone-induced reproductivecosts of activity. In male-larger species, costs of reproductionin terms of growth are suggested by supplemental feeding ofreproductive female C. atrox in their natural environment andby experimental manipulation of reproductive cost in femaleS. jarrovii. Growth costs of reproduction, however, do not contributesubstantially to the development of SSD in male-larger S. jarrovii.We conclude that the energetic costs of testosterone-induced,male reproductive behavior may contribute substantially to thedevelopment of SSD in some female-larger species. However, despitestrong evidence that reproductive investment exacts a substantialcost in growth, we do not support the reproductive cost hypothesisas a general explanation of SSD in male-larger species.     

The role of pineal and eyes in the photoperiodic effect on the gonad of the medaka, Oryzias latipes.

Studies on gonadal development in the female medaka, Oryzias latipes, intact, blinded or blinded-pinealectomized, maintained on different lighting schedules, i.e. LD14:10 and LD 10:14 and under natural daylight conditions, revealed that: 1)blinded fish were capable of responding to annual photoperiodic shifts with alterations of gonadal development, if the pineal was left intact; 2)the mean gonosomatic index(GSI) was higher in blinded fish than in intact fish, during a period from October through April, under LD 14:10, LD 10:14 and the natural daylight conditions; 3)when reared under the natural daylight conditions, the mean GSI in blinded fish was approximately half of that in intact ones in May and June, although the values were significantly larger than those in fish examined during all the other seasons; and 4)in December, the mean GSI was larger in blinded-pinealectomized fish than in intact and blinded ones. These findings seem to indicate that, during the breeding season, the gonadal growth in the medaka is stimulated by the pineal blocking the inhibitory mechanism against the hypothalamo-hypophyseal gonadotropic system, whereas during non-breeding seasons, the gonadal involution is induced by an anti-gonadal factor released by the pineal. It is likely that the effects of photoperiods are exerted on the pineal through the eyes.     

Seasonal changes in adrenal and gonadal activity in the quail, Perdicula asiatica: involvement of the pineal gland

The present study assessed annual adrenal gland activity in the Indian tropical Jungle bush quail, Perdicula asiatica. We also elucidated the role of the annual variations in gonadal steroids and melatonin in the regulation of its activity. Increasing day length (photoperiod), ambient temperature and rainfall are positively correlated with adrenal and gonadal functions, and inversely related to pineal gland activity. Pineal, adrenal and gonadal weights showed cyclical patterns relative to environmental factors, which were also correlated with plasma melatonin, corticosterone and gonadal steroids, respectively. In both sexes of P. asiatica, pineal gland weight and/or plasma melatonin levels were inversely related to adrenal lipids, (e.g. phospholipids, free and esterified cholesterol) and plasma corticosterone levels. Melatonin levels also showed an inverse relationship with plasma testosterone and estradiol levels. These studies indicate that changes in environmental factors promote annual variations in adrenal and gonadal activity probably by modulating the pineal gland. Melatonin receptors have been localized in the pars tuberalis, adrenal gland and gonads of birds, the pineal gland may, therefore, mediate environmental stimuli indirectly and directly to down regulate adrenal and gonadal activity, which run in parallel in this species.     

ASYNCHRONOUS REPRODUCTIVE ACTIVITY IN THE BROADCAST SPAWNER CELLANA CAPENSIS (GMELIN, 1791) (GASTROPODA: PATELLIDAE)

Short-term fluctuations in the reproductive condition of thelimpet Cellana capensis are described. Gravimetric and histologicalanalyses of gonadal development were carried out on samplescollected at 4 day intervals over a 3 month period from theMkambati Nature Reserve, Transkei. The results indicate a distinctlack of reproductive synchrony within the population, whichapplied not only to gametogenic development per se but alsoto spawning activity. Some of the spawning events identifiedduring the study period were more marked than others, thesemay reflect a greater degree of synchronization in activity.Results presented suggest that C. capensis is probably a ‘partial’rather than a ‘complete’ spawner. The implicationsof asynchronous spawning activity for the reproductive successof broadcast spawners are also discussed. (Received 6 December 1988; accepted 15 February 1989)     

Phenotypic variation in seasonal adjustments of testis size, body weight, and food intake in deer mice: role of pineal function and ambient temperature

We investigated pineal function as well as reproductive and energetic characteristics in male deer mice (Peromyscus maniculatus) that differentially respond to short photoperiod with full, partial or no gonadal regression. In mice at both high (23 °C) and low temperature (1 °C), these phenotypic differences in reproductive responses to short days were not reflected by differences in urinary excretion of 6-sulphatoxy-melatonin, the main metabolite of pineal melatonin. Neither duration nor amplitude or phase-angle of nocturnal peaks in 6-sulphatoxymelatonin significantly differed between reproductive phenotypes at either temperature. Differences in testis size were, however, associated with different energy requirements. In gonadally regressed males only, food intake and body weight were significantly (P < 0.01) reduced by up to 29% and 13% respectively. Chronic cold exposure (5 °C) had no effect on the proportion of males undergoing testicular regression under short days, but caused a general elevation in body weights among all mice (P < 0.05). Phenotypic differences in body weight and food intake were maintained in the cold. Together, these results suggest that within-population variation of reproductive responses in male deer mice is based on post-pineal differences in the regulation of gonadal function, and that phenotypic characteristics in reproductive and energetic responses to short days are largely unaffected by ambient temperature. Accepted: 2 October 1995     

Growth and Reproductive Strategy in VeronicaSpp.

Seed production, germination and plant growth form were studiedin several species of Veronica. The objectives were to comparethe reproductive strategies of annual and perennial speciesof Veronica which varied in their habitats and aggressivenessas weeds. ‘Reproductive effort’ was fairly similar for bothannuals and perennials, falling within the commonly observedrange for annual species. Seed number per capsule and the 100-seedweight were unaffected by the growth conditions but plant growthform was substantially modified in V. agrestis. The small-seeded Veronica species required light for germinationand the winter annual V. hederifolia germinated much more vigorouslyafter a low temperature treatment. The growth form, vigour andextent of adventitious root production and rapid seed germinationof V. persica must be significant factors in the spread andestablishment of this species. Veronica spp., germination, growth form, reproductive strategy     

Morphogenesis of Asymmetry and Symmetry in Lemna perpusilla Torr

The determination of which of the two reproductive pockets ofLemna perpusilla will produce the first daughter frond undernon-flowering conditions, or the flower under inductive conditions,is related to asymmetric gradients within the mother frond whichmay be the result of the asymmetric position of the axillaryfrond. Treatments that encourage vigorous growth of axillaryfronds may overcome the normal situation of asymmetry and causea morphological symmetry which is qualitatively different fromthe symmetry induced by treatments (TIBA) which prevent thedevelopment of axillary fronds. The origin and maintenance ofasymmetry in Lemna are discussed. Lemna perpusilla Torr., morphogenesis, asymmetry of fronds, tri-iodobenzoic acid     

Variation in reproductive success and gonadal allocation in the simultaneous hermaphrodite,Serranus fasciatus

Summary This paper examines the correlates of individual size, reproductive success, gonadal allocation, and growth in a hermaphroditic reef fish. Individuals in S. fasciatus mature as simultaneous hermaphrodites; large individuals subsequently lose female function and become functional males. Daily female reproductive success was highly correlated with both hermaphrodite size and amount of female gonadal tissue. Three separate comparisons gave a positive correlation between male reproductive success and male gonadal allocation: (1) Males had higher levels of male gonadal allocation and male reproductive success than hermaphrodites. (2) The percent of gonad allocated to male tissue in hermaphrodites was higher in the year they had higher male mating success. (3) Male gonadal tissue of hermaphrodites was positively correlated with male reproductive success in the year that male reproductive success by hermaphrodites was higher and more variable. There was no evidence for a trade-off between male function, female function, and growth among hermaphrodites. Many of these patterns have also been observed in plants, but the selective pressures leading to these patterns in S. fasciatus and plants are probably quite different.     

Biological Nitrogen Fixation in Pachyrhizus ahipa (Wedd.) Parodi

The patterns of plant growth and N₂ fixation capability in Pachyrhizusahipa (Wedd) Parodi inoculated with Bradyrhizobium ‘PachyrhizusSpec 1’ strains (Lipha Tech) were investigated in a zero-Nculture system under greenhouse conditions The P ahipa plantis day-neutral with respect to reproductive development Competitionoccurred between the two storage organs (legume and tuber) andprevented high tuber yield in P ahipa The symbiotic effectivenessof the association was high, as the profuse nodulation providedthe inoculated plants with adequate amounts of N Nodules werepresent throughout the cycle of P ahipa The change in rate ofN₂ fixation (RNF) and relative growth rate (RGR) was almostparallel during ontogenesis The developmental pattern of N₂fixation activity revealed that 65% of total N₂ fixation occurredafter N began to accumulate in the reproductive (pod wall plusseed) tissue During pod filling allocation of N compounds tothe seeds exceeded N₂ fixation, the pod walls being the primarysource of redistributed N, followed by the leaves. Pachyrhizus ahipa (Wedd) Parodi, ahipa, tuber crop, dinitrogen fixation, dry matter, N partitioning, reproductive growth     

Same sexual system but variable sociobiology: evolution of protandric simultaneous hermaphroditism in Lysmata shrimps

The sexual system of the decapod (caridean) shrimp Lysmata isprotandric simultaneous hermaphroditism (PSH). Individuals firstmature as males (male phase = MP) and then when older (larger)change to the external phenotype of female carideans (femalephase = FP). However, unlike purely protandric carideans, LysmataFPs retain reduced male gonadal tissues and ducts, and are ableto mate nonreciprocally as males as well as to reproduce asfemales. Thus, FPs of Lysmata species are functional simultaneoushermaphrodites although most reproductive effort is devotedto embryo production and incubation. The question explored hereis, given the propensity of carideans to protandry, the apparentlow cost, and high reproductive advantage of PSH, why has notPSH evolved more frequently? The mating systems and sexual selectionof caridean shrimps, the original sex of protandric individuals,the cost of maleness, and sex allocation theory are discussedin relation to protandry and PSH. None of these factors adequatelyexplains the evolution of PSH of Lysmata species. Lysmata hasat least 2 species groups with very different sociobiologies;these groups do not appear to share current selective pressuresthat would explain PSH in both. A historical contingency hypothesis,testable in part with a phylogenetic analysis, may explain theevolution of PSH in Lysmata.     

How outcrossing hermaphrodites sense the presence of conspecifics and suppress female allocation

One of the greatest remaining problems in the field of sex allocationis the variation across species of the extent to which individualsadjust their offspring sex ratio or sex allocation accordingto environmental conditions. Particularly in the field of sexallocation, there is a lack of research on the mechanisms andcues involved. We address this problem in the outcrossing simultaneoushermaphroditic polychaete Ophryotrocha diadema, which shiftssex allocation by suppressing the female function in the presenceof reproductive competitors or multiple partners. We documentthat O. diadema hermaphrodites can evaluate group size by meansof a species-specific chemical released by mature individualsin water. The perception of such a water-borne signal is sufficientto trigger the observed shift in sex allocation, i.e., a suppressionof the female function, irrespective of the frequency of encounterswith conspecifics. Under natural conditions, such a pheromonewill favor partners' encounters and the adjustment of individualsex allocation according to the numbers of reproductive competitorsand/or potential partners.     

THE ANATOMY OF CYLINDROTIS QUADRASI FORM OKINAWA ISLAND, JAPAN AND THE SUBFAMILIAL POSITION OF THE GENUS CYLINDROTIS MOLLENDORFF, 1895 (ARCHAEOPULMONATA: ELLOBIIDAE)

Cylindrotis quadrasi Mollendorff, 1895, the type species ofthe ellobiid genus Cylindrotis, was collected from Okinawa Island,southwest Japan. This is the first record of C. quadrasi inJapan and the third in the world. The anatomical charactersof this species, i.e. shell, head-foot, pallial cavity, radula,stomach, reproductive system, and central nervous system, aredescribed in detail for the first time. The significance ofthese observations is discussed and the subfamilial positionof Cylindrotis is reconsidered. Cylindrotis was included inthe Pythiinae, the most primitive group of the Ellobiidae, basedon the conchological characters alone. However, the male andfemale tracts of the reproductive system are entirely separated,and other internal organs exhibit highly derived conditions.Accordingly, Cylindrotis is transferred from the Pythiinae tothe Ellobiinae. Among the genera of the latter subfamily, Cylindrotisseems to be allied to Blauneria Shuttleworth, 1854 and LcucophytiaWinckworth, 1949. (Received 1 February 1993; accepted 7 June 1993)     

BUCEPHALID TREMATODE INFECTIONS IN MYTILID BIVALVES FROM THE ROCKY INTERTIDAL OF SOUTHERN CHILE

Bucephalid trematode sporocysts were found embedded amidst themantle tissue and viseral mass of intertidal mytilid bivalvesfrom Southern Chile. An investigation into the prevalence ofinfection revealed that <1% of Perumytilus purpuratus wereinfected whereas 20–32% of the Semimytilus algosus examinedharboured sporocysts. No sporocysts were found in an adjacentpopulation of Choromytilus chorus. Prevalence of infection wasrelated to size of S. algosus present at a particular site,and generally increased with host size. The majority of parasitizedmussels appeared to be at an advanced stage of infection, inthese animals destruction of the host's gonadal tissue was evident.Such parasitic castration combined with high prevalence of infectionsuggests that the reproductive potential of this mytilid maybe severely impaired. The implications of these results forthe mariculture industry are also discussed. (Received 19 February 1991; accepted 11 June 1991)     

Seasonal effects of pinealectomy on testicular recrudescence and secretory activity of seminal vesicles in the catfish Heteropneustes fossilis (Bloch)

The effects of pinealectomy on testicular activity and secretory activity of seminal vesicles were examined in the catfish Heteropneustes fossilis under various combinations of photoperiod and temperature during different periods of the annual reproductive cycle. Pinealectomy had no effect on gonadal activity during the preparatory, prespawning and spawning periods of the reproductive cycle. However, during the postspawning period, under long (LD 14:10) or short (LD 9:15) photoperiod at 25° C or at gradually increasing ambient temperature, pinealectomy accelerated testicular recrudescence and secretory activity of the seminal vesicles. Nevertheless. during this period the presence of the pineal facilitated the recrudescence of testes and seminal vesicles in catfish exposed to continuous light (LL), continuous darkness (DD) and 12 hL:l2 hD (LD) at 25° C. These findings suggest that the role of the pineal in catfish reproduction is variable and depends upon the photoperiod and temperature regimes to which the fish are exposed, as well as on the time of the year and the state of the reproductive cycle. The results also suggest that the effects of pinealectomy in catfish are mediated through an influence on the hypothalamo-hypophyseal gonadal axis.     

Physiological Aspects of Zebra Mussel Reproduction: Maturation, Spawning, and Fertilization

SYNOPSIS. The prolific reproductive capabilities of the zebramussel, Dreissena polymorpha, have facilitated the rapid spreadand high densities of this biofouling organism since its accidentalintroduction into North America less than 10 years ago. Researchon its reproductive mechanisms and capabilities may be valuablenot only in predicting its further spread, but also in investigatingbasic mechanisms of reproduction and development and in developingnew strategies to mitigate its impact. Since zebra mussels aredioecious and fertilization occurs externally, coordinated maturation,spawning, and other mechanisms have evolved to increase theprobability of successful fertilization. The zebra mussel undergoesan annual cycle of gonadal growth and gamete maturation, culminatingin one or more spawning events in late spring or early summer.Temperature, rates of temperature change, food availability,and effects of neighboring mussels seem to be critical variablesthat determine reproductive responses. Serotonin is a biogenicamine which is implicated in spawning behavior and can reliablytrigger spawning. Serotonin is present in the gonad in neuralvaricosities that encircle groups of gametes, and specific serotonergicligands can mimic or block spawning caused by serotonin. Infemales, serotonin reinitiates meiosis causing maturation fromprophase I to metaphase I prior to spawning. Spawned oocytescontain substances that are species specific sperm chemoattractants.The sequence of binding, entry, and subsequent nuclear movementshave been observed with fluorescence and scanning microscopy.Despite their negative ecological and economic impacts, zebramussels have also provided a new and easily obtainable resourcefor studies of reproductive mechanisms.     

PLASTICITY OF THE LIFE CYCLE OF XEROPICTA DERBENTINA (KRYNICKI, 1836), A RECENTLY INTRODUCED SNAIL IN MEDITERRANEAN FRANCE

Xeropicta derbentina (Krynicki, 1836), a native of Eastern MediterraneanEurope, was introduced to southeastern France during the 1940sand is now widely spread across Provence. In summer it aggregateson plants, making its populations clearly visible. However,its life cycle within the Mediterranean basin is poorly documented.While X. derbentina in its native area exhibits an annual lifecycle, this species has been found in Provence to have a bienniallife cycle. Moreover, in southeastern France, field studieswithin a restricted area show variations in demographic structure.In consequence, the life cycle of X. derbentina and the demographicpatterns observed require clarification. Five populations withvarious demographic structures were studied over 1 year in thesame location, i.e. under the same climatic conditions. Thefield study was complemented by laboratory observations on mating,egg-laying and hatching. Xeropicta derbentina appears to bea semelparous species, with an annual life cycle being foundon four plots. The reproductive period begins at the end ofsummer and lasts until the beginning of winter. First egg-layingoccurs within 1 week after mating and lasts up to 30 days. Hatchingtakes place 15–20 days after egg-laying. Xeropicta derbentinapossesses multiple mating and egg-laying sessions, involvingsuccessive hatching. Populations are mainly characterized bytwo growth stages, the first in spring when newly-hatched snailsevolve into juveniles, and the second in late summer when theyreach maturity. However, on the highest density plot, a bienniallife cycle is observed for some newly-hatched snails that showan interrupted growth during summer and evolve into juvenilesonly in the second autumn. Moreover, this life cycle not onlyvaries among plots but also at a 1-year interval within plots.Hence, the life span of X. derbentina is between 12 and 20 months,but can be extended up to 30 months according to whether hatchingoccurs early or late and whether they survive the first andsecond winters. Xeropicta derbentina is thus able to have variousgrowth speeds and life spans, and appears to switch from anannual life cycle to a biennial cycle in response to populationdensity or climatic conditions. (Received 8 October 2004; accepted 15 December 2004)     

The Comparative Reproductive Physiology of Sea Turtles

Although the reproductive ecology of marine turtles has beenthe subject of numerous long-range studies, the reproductivephysiology of these unique animals is little known. Recently,however, preliminary anatomical and endocrinological studieshave provided a good basis on which to begin to attempt to explainseveral of their unusual biological systems. New findings relatedto the anatomy of Chelonia mydas and Lepidochelys olivacea arepresented. The female, although having a very massive pair ofsimultaneously functioning ovaries, appears in most ways verysimilar to other chelonians. Corpora hemorrhagica, corpora luteaand corpora atretica from active ovaries are briefly described.Ovulation coincides with a luteinizing hormone and progesteronesurge. Sperm are probably stored for the season after a singlemating period which appears to occur prior to the first ovulation.Males may also cycle and mating as well as nesting are seenas more or less seasonal. The seasonality could be controlledin part by melatonin or other endocrines from the sea turtle'smassive pineal complex. A hypothetical model for reproductionis presented in hopes of stimulating interest in physiologicalapproaches to the study of marine turtle reproduction.     

Pineal-dependent and -independent effects of photoperiod on immune function in Siberian hamsters (Phodopus sungorus)

Siberian hamsters (Phodopus sungorus) exhibit reproductive and immunological responses to photoperiod. Short (<10-h light/day) days induce gonadal atrophy, increase leukocyte concentrations, and attenuate thermoregulatory and behavioral responses to infection. Whereas hamster reproductive responses to photoperiod are dependent on pineal melatonin secretion, the role of the pineal in short-day induced changes in immune function is not fully understood. To examine this, adult hamsters were pinealectomized (PINx) or sham-PINx, and transferred to short days (9-h light/day; SD) or kept in their natal long-day (15-h light/day; LD) photoperiod. Intact and PINx hamsters housed in LD maintained large testes over the next 12 weeks; sham-PINx hamsters exhibited gonadal regression in SD, and PINx abolished this effect. Among pineal-intact hamsters, blood samples revealed increases in leukocyte, lymphocyte, CD62L+ lymphocyte, and T cell counts in SD relative to LD; PINx did not affect leukocyte numbers in LD hamsters, but abolished the SD increase in these measures. Hamsters were then treated with bacterial lipopolysaccharide (LPS), which induced thermoregulatory (fever), behavioral (anorexia, reductions in nest building), and somatic (weight loss) sickness responses in all groups. Among pineal-intact hamsters, febrile and behavioral responses to LPS were attenuated in SD relative to LD. PINx did not affect sickness responses to LPS in LD hamsters, but abolished the ameliorating effects of SD on behavioral responses to LPS. Surprisingly, PINx failed to abolish the effect of SD on fever. In common with the reproductive system, PINx induces the LD phenotype in most aspects of the immune system. The pineal gland is required for photoperiodic regulation of circulating leukocytes and neural-immune interactions that mediate select aspects of sickness behaviors.     

The Effects of Infection by Rust (Puccinia lagenophorae Cooke) on the Growth of Groundsel (Senecio vulgaris L.) Cultivated under a Range of Nutrient Concentrations

Groundsel (Senecio vulgaris L.), healthy or infected with therust fungus Puccinia lagenophorae Cooke, was grown at a rangeof nutrient concentrations in sand culture. There were statisticallysignificant interactions between the effects of infection andnutrient supply upon the dry weights of stems, leaves, rootsand reproductive tissues, leaf area and cumulative capitulumproduction. This interaction occurred since infection causedsignificant inhibitions of growth only at moderate or high nutrientconcentrations. At low concentrations rusted plants were similarto or slightly larger than controls. Both in controls and rustedplants root: shoot ratios increased as nutrient supply declined.The ratio of root: shoot dry weight was consistently reducedby infection whilst root length: leaf area ratio was relativelyunchanged. More detailed investigations confirmed that infection had littleeffect on plant growth under nutrient deficient conditions despitesuppression of the host's ability to increase root: shoot ratiosin response to nutrient stress. This reflected the inhibitionof relative growth rates in rusted plants at high but not lownutrient concentrations, which in turn reflected reduced netassimilation rates (NAR). Increases in leaf-area ratio (LAR)often ameliorated the decline in NAR in rusted plants. Senecio vulgaris L., Puccinia lagenophorae Cooke, nutrient deficiency, growth, root: shoot ratio