Variation in plasma lipids during the reproductive cycle of male and female desert tortoises,Gopherus agassizii

Plasma triacylglycerol, phospholipid, cholesterol, cholesterol esters, fatty acids, and total lipids were measured in 30 female and 20 male desert tortoises (Gopherus agassizii) during the annual reproductive cycle in the eastern Mojave desert, Nevada. Blood samples were collected at monthly intervals from April to October. All lipid fractions, with the exception of free fatty acids, were significantly higher in female plasma than in male plasma in all months of the year. In contrast, free fatty acids were higher in male plasma than in female plasma in all months. The seasonal pattern in estradiol secretion mirrored that of triacylglycerol, phospholipid, cholesterol, and total lipid, all of which showed a significant correlation with the hormone. Estradiol and the vitellogenesis-associated lipids were all significantly higher in August, September, October, and April than in June. The seasonal variation in cholesterol ester levels in females did not correlate with any of the reproductive events and did not appear to be involved in yolk precursor formation. Total lipid in males showed a negative correlation with testosterone and spermatogenesis. Individual fatty acids in the June and August samples (at the highest and lowest estradiol levels) were compared in male and female plasma. The percent of C18:3n3, C18:2n6, C18:1n9, C20:5n3, and C22:5 were significantly higher in the June female plasma sample than in the August sample. Docosahexanoic (C22:6n3) acid was barely detectable in female plasma in either month.     

Effects of climatic variation on field metabolism and water relations of desert tortoises

We used the doubly labeled water method to measure the field metabolic rates (FMRs, in kJ kg^?1?day^?1) and water flux rates (WIRs, in ml H₂O?kg^?1?day^?1) of adult desert tortoises (Gopherus agassizii) in three parts of the Mojave Desert in California over a 3.5-year period, in order to develop insights into the physiological responses of this threatened species to climate variation among sites and years. FMR, WIR, and the water economy index (WEI, in ml H₂O?kJ^?1, an indicator of drinking of free water) differed extensively among seasons, among study sites, between sexes, and among years. In high-rainfall years, males had higher FMRs than females. Average daily rates of energy and water use by desert tortoises were extraordinarily variable: 28-fold differences in FMR and 237-fold differences in WIR were measured. Some of this variation was due to seasonal conditions, with rates being low during cold winter months and higher in the warm seasons. However, much of the variation was due to responses to year-to-year variation in rainfall. Annual spring peaks in FMR and WIR were higher in wet years than in drought years. Site differences in seasonal patterns were apparently due to geographic differences in rainfall patterns (more summer rain at eastern Mojave sites). In spring 1992, during an El Niño (ENSO) event, the WEI was greater than the maximal value obtainable from consuming succulent vegetation, indicating copious drinking of rainwater at that time. The physiological and behavioral flexibility of desert tortoises, evident in individuals living at all three study sites, appears central to their ability to survive droughts and benefit from periods of resource abundance. The strong effects of the El Niño (ENSO) weather pattern on tortoise physiology, reproduction, and survival elucidated in this and other studies suggest that local manifestations of global climate events could have a long-term influence on the tortoise populations in the Mojave Desert.     

Reproductive effort and costs of reproduction in female European ground squirrels

Reproductive effort, factors affecting reproductive output and costs of reproduction were studied in primiparous yearling compared to multiparous older female European ground squirrels (Spermophilus citellus). Yearling females weaned smaller litters than older ones. Litter size increased with posthibernation body mass at the expense of slightly lighter young for yearling but not for older mothers. In older females, on the other hand, emergence body mass influenced offspring mass, whereas litter size was affected by oestrus date. High reproductive effort entailed reproductive costs in terms of reduced subsequent fecundity but not subsequent survival for both yearling and older females. The production of large litters and long duration of lactation delayed subsequent oestrus, which, in turn, correlated negatively with litter size. During the second half of lactation, oestradiol levels were significantly elevated, indicating the initiation of follicular maturation processes. Oestradiol levels during that time correlated negatively with current, but positively with subsequent litter size. We therefore assume that inhibitory effects of lactation on gonadal development may mediate the negative relationship between reproductive effort and subsequent reproductive timing in adults. This effect is absent in yearlings because they are reproducing for the first time. Reproductive output in yearlings was influenced by interactions between structural growth and puberty. Received: 22 March 1999 / Accepted: 7 June 1999     

Reproductive effort of female bank voles in a risky environment

In cyclically fluctuating vole populations, strong intraspecific competition and intense predation simultaneously or separately increase the costs of reproduction and so may set the framework for the optimal breeding tactic of voles. In a factorial experiment, we manipulated two factors in the breeding environment of bank vole (Clethrionomys glareolus) pairs, each with two treatment levels: no predation risk or high risk of specialist predators and low or high density of voles. In the manipulation, we used odours of conspecific voles and/or mustelid predators. Both over-wintered and young, summer-born, wild-caught bank vole females and males in breeding condition were used. Each female—male pair of voles was placed in a cage and the cages were distributed in large outdoor enclosures. All animals were fed ad libitum. Under predation risk, both old and young females suppressed breeding significantly. The density of conspecific voles did not affect overall breeding. However, there was some evidence that population density stimulated breeding of old females but suppressed breeding of young ones. Both risk factors appeared to increase litter sizes of those individuals who decided to breed. Our results indicate that the risk of predation may be an important factor determining reproductive tactics of bank vole females. In risky environments, females seemed to choose between two totally opposite tactics: they suppressed breeding, which may increase their own survival to the next breeding event, or they continued to breed in spite of expected high survival costs. Females seemed to compensate the latter costs with a higher effort to the current and probably the last reproduction.     

Subject: Reproductive effort and success

Research Notes on Avian Biology 1994: Selected Contributions from the 21st International Ornithological CongressGeneral Biology: Reproductive Success

Subject: Reproductive effort and success     

Sex differences in plasma corticosterone in desert tortoises, Gopherus agassizii, during the reproductive cycle

Blood samples from 30 female and 20 male adult desert tortoises, Gopherus agassizii, were collected at monthly intervals during the annual reproductive cycle (April to October). Plasma corticosterone and the sex steroids in each of the samples were analyzed by radioimmunoassay. Mean corticosterone levels in males were significantly higher than in females (P < 0.001) in every month. Male tortoises showed a marked seasonal pattern in plasma corticosterone with a highly significant peak in July, August, September, and October that corresponded with a similar peak in plasma testosterone. Testosterone and corticosterone in the male showed a highly significant correlation (P < 0.0001). The pattern of corticosterone in the female was less marked, with a significant peak in May during the mating and nesting season, but no association with the peak in estradiol in late summer was apparent. The highest levels of corticosterone in the males were associated with the peak in spermatogenesis and intense male-male combat. These results support similar data from other reptiles that suggest increased glucocorticoid secretion during periods of increased activity and metabolism.     

Reproductive study of Giant tortoises on Aldabra

A population of 150,000 Giant tortoises occurs on the Indian Ocean atoll of Aldabra. During the course of an extensive census and marking programme a preliminary reproductive study was carried out. This paper describes the pattern of seasonal ovarian development and the seasonal cycle of breeding activity that was observed.     

Mycoplasmosis in free-ranging desert tortoises in Utah and Arizona

Upper respiratory tract disease (URTD) has been associated with major losses of free-ranging desert tortoises (Gopherus agassizii) in the southwestern United States. This prompted a clinical examination of 63 free-ranging desert tortoises for signs of URTD and sampling for Mycoplasma agassizii, the causative agent of URTD. Tortoises were sampled from three sites in the eastern Mojave Desert (1992-93), and from three sites in the Sonoran Desert (1992-94). Plasma samples were tested for antibodies to M. agassizii using an enzyme-linked immunosorbent assay (ELISA). Nasal aspirate samples from 12 Sonoran tortoises were tested using polymerase chain reaction (PCR) test directed at the 16S rRNA gene of M. agassizii. Nasal aspirate samples from all tortoises were cultured for M. agassizii. In the Mojave Desert, nine tortoises had clinical signs of URTD and eight were seropositive for M. agassizii. In the Sonoran Desert, there were no clinical signs of URTD, but two tortoises were seropositive, and two tortoises had positive PCR results.     

Paternal investment directly affects female reproductive effort in an insect

Female reproductive effort can be influenced by the quality of her mate. In some species, females increase their reproductive effort by differentially allocating resources after mating with high-quality males. Examination of female reproductive effort in relation to male quality has implications for estimating the evolvability of traits and for sexual-selection models. Accurate quantification of reproductive investment is not possible in many species. Butterflies are an exception, as most nectar-feeding species emerge with almost intact reproductive resources, and in some species males provide nutrients at mating that enhance female fecundity. By manipulating male donations and using radioactive isotopes, we quantified the effect of variation in nutrient provisioning on female reproductive effort in two butterfly species. In the greenveined white butterfly, Pieris napi, females increased their reproductive effort after receiving large male donations. By contrast, in the speckled wood, Pararge aegeria, where males do not provide nutrients, female reproductive effort was independent of male ejaculate. Increased reproductive effort in Pieris napi resulted from the production of more eggs, rather than from investing more resources per egg. In this species donating ability is heritable; hence females laying more eggs after mating with high-donating males benefit both through higher fecundity and through the production of high-donating sons.     

Parental effort and reproductive skew in coalitions of brood rearing female common eiders

Members of breeding groups face conflicts over parental effort when balancing antipredatory vigilance and feeding. Empirical evidence has shown disparate responses to manipulations of parental effort. We develop a model in which we determine the evolutionarily stable effort of partners given their body conditions, allowing the benefits of shared care to be unevenly divided, and we test this model's predictions with data on common eiders (Somateria mollissima). Eiders show uniparental female care; females may share brood rearing, or they may tend alone, and their body condition at hatching of the young shows large environmentally induced variation. The model predicts that parental effort (vigilance) in a coalition is lower than when tending alone, controlling for parental condition; this prediction is supported by the data. The parental effort in a coalition should be positively correlated with body condition, and this prediction is also supported. Finally, parental effort should increase when partner condition decreases and vice versa; this prediction is partially supported. The Nash bargaining game may provide promising avenues by which to determine the precise settlement of reproductive skew and effort between coalition partners in the future.     

Energetics of age-specific reproductive effort in female Harp seals, Phoca groenlandica

The theory of reproductive effort is commonly viewed in the perspective of the cost of offspring to the parent. This interpretation predicts that older females make a greater reproductive effort than younger females, for whom the cost of lost future offspring is less. The benefit early-born offspring represent in future generations is greater than that of late-born offspring in most natural populations, so, alternatively, young females should make a greater reproductive effort. These hypotheses were investigated in nursing Harp seals by examining age-specific reproductive effort in terms of energetic maternal investment.
Younger Harp seal females lost weight at the same rate as did older females. Moreover. since younger, growing females tended to start lactation with less blubber than older females. weight loss during 10 days (d) of lactation represented a greater proportion of stored energy for young females. A successfully weaned pup, therefore, constituted a greater reproductive effort for them than for older females, consistent with benefit–oriented predictions from theory.
A model is presented to outline the components of an annual energy budget for female Harp seals in the context of reproductive effort. This model indicates that young females may devote a greater proportion of their annual net energy to reproduction and take a greater risk of lost future reproduction. It also identifies areas of inadequacy of existing data on pinniped energy budgets.     

Patterns of reproductive effort in male and female shrubs of Oemleria cerasiformis: a 6-year study

1 We monitored flowering and fruiting of individual male and female plants of Oemleria cerasiformis over a 6-year period in a population in western Canada, and calculated fruit set (percentage of pistils maturing) and reproductive effort (RE) (gram of reproductive tissue per gram of leaf).
2 Over 6 years, male O. cerasiformis had on average much lower total RE, but much higher RE at flowering, than females.
3 In males, strong correlations between RE and light suggested that investment in reproduction was largely determined by light levels. There were strong positive correlations of RE between years, with no evidence of periodic fluctuations.
4 In females, in contrast to males, RE at flowering was not related to light. However, fruit set was strongly correlated with light. Flowering RE and fruit set were uncorrelated in females, indicating that these are affected by different factors.
5 Correlations of RE between years in females, although often significant, were lower than in males, indicating that RE fluctuates more between years in females than in males and may respond to past levels of RE. Flowering may reflect adjustments in response to past reproduction, or may be controlled by resources other than light. Fruit set was not significantly related to previous RE.
6 The greater total RE of females and their limited ability to adjust fruit set are probably major factors contributing to the greater mortality rates of females and the male-biased sex ratios in O. cerasiformis .     

Reproductive effort in desert versus mediterranean crucifers: the allogamous Erucaria rostrata and E. hispanica and the autogamous Erophila minima

Components of the total sexual investment of plants growing under mediterranean climatic and edaphic conditions were compared with those of plants growing in the desert, in the closely related allogamous species pair Erucaria hispanica and E. rostrata and in populations of the autogamous species Erophila minima. In all cases lower total investment was evident in the desert plants. At the prezygotic phase it was expressed by (1) reduced production of flowers; (2) lower allocation to the production of male gametophytes and some floral organs; and (3) packaging of more ovules per ovary. The ratio of reproductive: vegetative biomass which was found to be greater in the desert plants and their lower pollen: ovule ratio are perhaps indicative of greater efficiency. Their smaller investment at the postzygotic phase was expressed in: (1) reduction in total numbers of fruits and seeds; (2) decrease in seed size and weight. Yet in the desert plants the number of seeds per total biomass was found to be significantly larger and fertility rates (seed-set per ovule, fruit-set per flower per plant) were equal to or greater than those in the mediterranean plants. The trends observed in this study in desert plants, which may result in more efficient exploitation of resources, are similar in the species involved, regardless of their breeding system-autogamous or allogamous.     

Reduced reproductive effort in male field crickets infested with parasitoid fly larvae

Some populations of the field cricket Teleogryllus oceanicusare parasitized by the phonotactic fly Ormia ochracea. Flieslocate crickets by their song and deposit larvae onto them.The larvae develop inside the cricket for 1 week before killingthe host upon emergence. The reproductive compensation hypothesispredicts that parasitized crickets should increase their reproductiveeffort during the initial stages of infestation to offset theloss of fitness resulting from their shortened life span. An alternative hypothesis predicts that parasitized crickets willdecrease reproduction, either because they are unable to reproduceor because selection acting on the parasitoid favors decreasedhost reproduction. In laboratory experiments, parasitized malecrickets had reduced reproductive effort (spermatophore production,calling, mating activity, and mass allocated to reproductivetissue) compared to unparasitized males. Parasitized males fedad libitum showed no evidence of allocating a greater proportionof their resources to reproduction. Parasitized and healthymales did not differ significantly in resting or maximal metabolicrates, although this may have been due to the substantial contributionof larval respiration to the metabolic rate of the host—parasitoidcomplex. These results are consistent with previous studiesand suggest that T. oceanicus males parasitized by O. ochraceado not increase their reproductive effort. We discuss potentialreasons that crickets do not increase reproductive effort inresponse to fly larvae and address difficulties in demonstratingaltered life-history patterns in response to parasitism.     

The timing of male reproductive effort relative to female ovulation in a capital breeder

1. In large herbivores, the timing of breeding is important for females to hit peak plant protein levels. For males, the timing of reproductive effort is important to maximize the number of females they can mate during autumn rut in competition with other males. The latter depends on when most females are ovulating, but also on how other males with a different competitive ability are timing use of their capital (fat); it may pay younger males to invest more heavily later when prime aged males are exhausted. 2. Based on estimates of body mass loss, we quantify how much timing (start, peak and end dates) of male reproductive effort during rutting varies depending on male age, density and climate as well as timing of female ovulation. 3. Ovulation in adult females was delayed by 5 days from low to high density, and ovulation was also more synchronous at high density. The starting date of decline in male body mass was only later in yearlings than among other age groups. However, at low density, peak and end dates of rut became increasingly earlier and close to peak female ovulation with increasing age up to 7 years of age. Prime-aged males matched peak effort closely with peak rate of prime-aged female ovulation, while younger males were delayed. This is consistent with the view that younger males have a better chance when the prime-aged males are becoming exhausted. 4. Apart from yearlings, male age groups were synchronized in both the starting, peak and end dates of mass decline at high density. Thus, this partly follows change in female ovulation patterns, but also suggests that competition among males decreased with increasing density due probably to lower intensity of sexual selection. The lowered sexual selection may be due not only to more synchronous female ovulation, but also increasingly female-biased sex ratios and a younger male age structure with increasing density. 5. The onset of rutting is somewhat independent of male age (apart from the youngest males), but the peak and end of rutting effort is dependent strongly upon age, density and peak female ovulation. Male rutting phenology is thus best interpreted as a compromise between hitting peak female ovulation and intensity of sexual selection.     

Reproductive effort in invasive and non-invasive Rubus

We quantified the physiological costs and the total amount of resources allocated to reproduction in two closely related species of Rubus, one of which is invasive. These two species share several morphological and life-history characteristics and grow together in the Pacific Northwestern United States. Reproductive effort was manipulated in canes of both species by removing flower buds. The non-invasive species, R. ursinus, exhibited significantly greater water stress in the reproductive canes, as indicated by lower leaf water potential (O) and reduced stomatal conductance (g_s). This species also showed a reduction in leaf nitrogen concentration ([N]) associated with reproduction. Combined, these factors led to reduced photosynthesis (A) on a diurnal basis, lower water-use efficiency as inferred from '¹³C, and reduced photosynthetic capacity. All of these effects were more pronounced during the fruiting stage than in the flowering stage. The invasive species, R. discolor, showed no changes in water stress, [N], '¹³C, or A associated with reproduction. A model was used to estimate total gross photosynthesis (A_gross) for reproductive and non-reproductive canes of both species over cane lifetime. Reproduction was associated with a greater decline in A_gross for the non-invasive R. ursinus than for the invasive R. discolor. Although R. discolor allocated more resources directly to flowers and fruit than R. ursinus, the invasive species had significantly lower reproductive effort, or total amount of resources diverted from vegetative activity to reproduction, than the non-invasive species. By minimizing the reduction of photosynthesis associated with reproduction, this invasive species may be able to minimize the trade-offs commonly associated with reproduction.     

Reproductive hacking: A male seminal protein acts through intact reproductive pathways in female Drosophila

Seminal proteins are critical for reproductive success in all animals that have been studied. Although seminal proteins have been identified in many taxa, and female reproductive responses to receipt of these proteins have been documented in several, little is understood about the mechanisms by which seminal proteins affect female reproductive physiology. To explore this topic, we investigated how a Drosophila seminal protein, ovulin, increases ovulation rate in mated females. Ovulation is a relatively simple physiological process, with known female regulators: previous studies have shown that ovulation rate is promoted by the neuromodulator octopamine (OA) in D. melanogaster and other insects. We found that ovulin stimulates ovulation by increasing OA signaling in the female. This finding supports a model in which a male seminal protein acts through “hacking” a well-conserved, regulatory system females use to adjust reproductive output, rather than acting downstream of female mechanisms of control or in parallel pathways altogether. We also discuss similarities between 2 forms of intersexual control of behavior through chemical communication: seminal proteins and pheromones.     

Nasal and cloacal bacteria in free-ranging desert tortoises from the western United States

Aerobic bacteria were collected from three free-ranging desert tortoise (Gopherus agassizii) populations in the eastern Mojave Desert (Arizona, Utah; USA) from 1989 to 1993, and from two free-ranging populations in the central Sonoran Desert (Arizona, USA) from 1990 to 1994. Six species of nasal bacteria and 18 species of cloacal bacteria were identified. At least one potential pathogen was found in the nasal cavity (Pasteurella testudinis), and at least two potential pathogens in the cloaca (Pseudomonas spp., Salmonella spp.).     

Hematology and plasma biochemistry reference range values for free-ranging desert tortoises in Arizona

Baseline values and ranges for 10 hematologic and 32 plasma chemistry parameters were analyzed for 36 free-ranging Sonoran desert tortoises (Gopherus agassizzi) collected in Yavapai and La Paz Counties (Arizona, USA) from 1990 to 1995. Tortoises were radio tagged from 1990 to 1994, and attempts were made to recapture them three times a year. Tortoises were weighed, measured, and chemically immobilized to collect blood for hematology and blood chemistry assessments. Tortoise biochemistry differed (P < 0.01) between sites and sexes and among seasons and years. Normal reference ranges for hematologic and plasma biochemistry parameters were determined. Seasonal and annual differences in hematology and blood chemistry were related to rainfall patterns, forage availability, and physiological condition.