Reproductive effort and floral photosynthesis in Spiranthes cernua (Orchidaceae)

This study examined the cost of reproduction and photosynthetic characteristics of the reproductive structures of Spiranthes cernua, an agamospermic, terrestrial orchid. Reproduction was frequent: two-thirds of the plants flowered at least 2 yr in a row and one-fourth of the consecutive-year runs were ~3 yr. Neither a significant decrease in leaf area nor a reduced likelihood of flowering was observed following 1 or 2 yr of inflorescence production. While there was a tendency for plants producing >16 flowers to have decreased size the next year, plants with the greatest number of flowers (31+) were the most likely to reproduce. Leaf and reproductive gas exchange were measured in the field. Low but positive rates of net photosynthesis were documented at all stages of inflorescence development. The average rates of photosynthesis for each stage were: leaves, 9.2 mmol CO2/m2s; inflorescence in bud, 3.7 mmol CO2/m2s; inflorescence in flower, 2.5 mmol CO2/m2s and infructescence, 0.2 mmol CO2/m2s. Based on diurnal gas exchange, the contribution of leaves and reproductive structures to seasonal carbon assimilation was 91.6 and 8.4%, respectively. The role of the inflorescence as a source and sink for carbon assimilation may lower the cost of reproduction and support frequent inflorescence production.     

Reproductive effort in molluscs

Summary A survey of the available molluscan literature shows that reproductive effort is higher in semelparous species (29.90%) than in iteroparous species (18.21%), and that in iteroparous species reproductive effort increases with successive breeding seasons. Oviparous species were found to divert considerably more into reproduction than viviparous species, with 24.24% and 5.25% channeled respectively.     

Reproductive effort in viscous populations

Abstract.— Here I study a kin selection model of reproductive effort, the allocation of resources to fecundity versus survival, in a patch-structured population. Breeding females remain in the same patch for life. Offspring have costly, partial long-distance dispersal and compete for breeding sites, which become vacant upon the death of previous occupants. The main result is that the evolutionarily stable reproductive effort decreases as offspring dispersal rate increases. The result can be understood as follows: In a well-mixed population with global competition, neither adults nor juveniles compete with relatives, but in a patch-structured population with dispersal restricted to the juvenile phase, juveniles experience relatively less competition with relatives than adults, thus making juveniles relatively more valuable. Because this asymmetry between adults and juveniles decreases with the dispersal rate, so does the evolutionarily stable level of allocation to fecundity.     

Reproductive effort reduces long-term immune function in breeding tree swallows (Tachycineta bicolor)

We examined whether strategies of reproductive allocation may reduce long-term immunocompetence through the effects of manipulated effort on secondary or acquired immunity. We tested whether increased reproductive effort leads to reduced immune function and survival by manipulating brood size in tree swallows (Tachycineta bicolor) and exposing breeding females to a primary and secondary exposure of sheep red blood cells to elicit a humoral immune response. Females raising enlarged broods produced fewer secondary antibodies than did females raising control or reduced broods. Most importantly, individuals with high secondary responses were more likely to survive to breed 3 years after brood manipulations, suggesting that differences in disease susceptibility may be caused by trade-offs in reproductive allocation. We also found that individual quality, measured by clutch initiation date, mediated the effects of brood manipulations, with higher-quality birds showing a greater ability to deal with increases in effort.     

Reproductive effort of Mastocarpus papillatus (Rhodophyta) along the California coast1

Species with sexual and asexual life cycles may exhibit intraspecific differences in reproductive effort. The spatial separation of sexual and asexual lineages, called geographic parthenogenesis, is common in plants, animals, and algae. Mastocarpus papillatus is a well‐documented case of geographic parthenogenesis in which sexuals dominate southern populations, asexuals dominate northern popula‐tions, whereas mixed populations occur throughout central California. We quantified abundances and reproductive effort of sexual and asexual fronds and tetrasporophytes at eight sites in California to test the hypotheses that (1) reduced sexual reproduction at higher latitudes and tidal heights explains the observed geographic parthenogenesis and (2) reproductive effort (spore production per blade area) declines with increasing latitude. Abundances of all phases varied site‐specifically. However, there was no geographic pattern of reproductive effort of fronds. Reproductive effort of fronds was greater in 2006 than in 2007 and correlated with sea surface temperatures. Sexual fronds exhibited greater reproductive effort than did asexual fronds and sexual reproductive effort was also inversely correlated with local upwelling index. Tetrasporophytes showed greater repro‐ductive effort in northern sites, but total supply of tetraspores per m² was greatest in the middle of the sampling range where crusts were more abundant. There was no decline of reproductive effort at higher latitudes. Geographic patterns of fecundity of life stages do not explain geographic parthe‐nogenesis in M. papillatus. Site‐specific differences in viability among spores or established thalli of different life cycles may explain their respective geographic distributions, as the sexual and asexual life cycles responded differently to environmental variations.     

Character displacement and coexistence in mud snails (Hydrobiidae)

Summary Populations of coexisting and of allopatrically occurring species of hydrobiid snails (Hydrobia ulvae, H. neglecta, H. ventrosa and Potamopyrgus jenkinsi) have been studied in 90 localities within three different areas. When H. ventrosa coexists with H. ulvae they show character displacement, i.e., the average body size of the former is smaller and that of the latter is larger. When these species live alone they are of approximately the same size. It is shown here that the size ratio between the coexisting species usually found (1.3–1.5) allows stable coexistence based on food praticles size selectivity alone. Variation in the degree of character displacement from locality to locality is explained by different degrees of genetical isolation of the populations. Coexisting H. ventrosa and H. ulvae have shorter, more well-defined periods of reproduction than they do when they occur alone. H. neglecta is larger than coexisting H. ventrosa and smaller than coexisting H. ulvae. Due to the patchy distribution and the fluctuating populations of this species, and due to the fact that pure H. neglecta populations are rare, data on this species are difficult to interpret. Potamopyrgus jenkinsi shows a different food particle size selection than the Hydrobia spp. of identical sizes. Competitive interactions between P. jenkinsi and coexisting Hydrobia spp. are therefore probably weak. In accordance with this, P. jenkinsi does not show character displacement when coexisting with, e.g., H. ventrosa.The fact that the major study area, the Limfjord, is only 150 years old as a marine habitat as well as various more recent man-made changes of the coast line allow estimates of the time scale of the microevolutionary changes which lead to character displacement.     

Reproductive effort decreases antibody responsiveness

The prevalence and intensity of parasitic infection often increases in animals when they are reproducing. This may be a consequence of increased rates of parasite transmission due to reproductive effort. Alternatively, endocrine changes associated with reproduction can lead to immunosuppression. Here we provide support for a third potential mechanism: reduced immunocompetence as a consequence of adaptive reallocation of resources in times of increased energetic demand. In captive zebra finches Taeniopygia guttata, reproductive effort was manipulated through brood size. Enhanced effort was found to affect the production of antibodies towards sheep red blood cells. In addition, activity of zebra finches was manipulated independently of parental care. Experimentally increased daily workloads in activity reward schedules also suppressed antibody production. Thus, we show that not just the reproductive state, but the increased activity that accompanies reproduction is associated with immunocompetence. This mechanism may be sufficient to explain the increased parasitism observed in reproducing animals. We suggest that reduced immunocompetence as a consequence of increased reproductive effort may be an important pathway for the life history cost of reproduction.     

Reproductive effort,fecundity and energy allocation in Marphysa sanguinea (Polychaeta: Eunicidae)

Summary

Reproductive effort in terms of fecundity and energy allocation was studied in the iteroparous and long lived polychaete Marphysa sanguinea. Both measures show great variability. Fecundity varied from 8500 to 24300 oocytes; no linear relationship was found between oocyte number and jaw length whereas a direct relationship was established between oocyte number and wet body weight. The energy content of germinal and somatic tissues was determined by differential scanning calorimeter (DSC). The reproductive effort for a single reproductive event was calculated according to the formula: RE = E_G/(E_G + E_S) where E_Gis the total energy of the germinal tissues and E_S is the total energy of the somatic tissues. The lack of correlation between reproductive effort and size index strongly suggests that reproductive allocation does not increase with age. The reproductive effort ranged from 0.04 to 0.19 with a mean value of 0.120.     

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 effort in cotton grass tussock tundra

Eriophorum vaginatum ssp. spissum is a dominant plant species of undisturbed cotton grass tussock tundra in Alaska. It also quickly invades and dominates recently disturbed sites. The hypothesis tested in this research was that the success of E. vaginatum on disturbed sites might be achieved through a higher allocation of biomass to reproductive structures relative to other tundra species. Reproductive allocation of tundra plants in general also was compared with plants of the temperate zone. The results indicate that E. vaginatum is about average among the common tundra species in terms of total reproductive allocation, allocation to seeds, and the proportion of total reproductive allocation that is accounted for by viable seeds. Tundra species, on a relative basis, allocate less biomass to all reproductive structures than temperate species but not necessarily less biomass to the output of viable seeds.     

Evidence for competition between mudsnails (Hydrodiidae): a field experiment

The paper describes an experimental investigation of competition between Hydrobia ulvae and H. ventrosa using enclosed populations at a site at which the species coexist naturally. lntraspecific competition is more intense than interspecific competition and may have a regulatory influence on snail densities. Other experimental studies of competition between mudsnails are reviewed and found to infer strongly the occurrence of competition in natural field populations. However, features of the species' field distributions previously attributed to interspecific competition can be more convincingly explained by other processes.     

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     

Reproductive effort and efficiency in the female common goby, Pomatoschistus microps (Krøyer) (Teleostei: Gobioidei)

Reproductive effort and efficiency are terms which together help to describe the reproductive investment of spawning fish. In this paper they are applied to a small and abundant inshore fish, the common goby, Pomatoschistus microps (Kroyer), based on data collected from the wild, and by modelling likely breeding patterns. Wet calorimetry was used to determine the energy content offish tissue, and this allowed reproductive effort to be expressed as both an energy and a weight ratio of gonad to somatic body. The gonadosomatic index (GSI) calculated as an energy ratio was greater than that calculated as a dry "weight ratio". For ripe females the GSI (dry weight) at the onset of breeding was 67.5%. Theoretical and observed values of the maximum number of egg batches laid by a single female during a 16-week breeding season are 9-10, and this corresponds to a total caloric investment of almost 12 kJ. Estimates of the reproductive efficiency, using the dietary energy content during the breeding season, range from 28.2% to a maximum of 44.0%. These values are among the highest calculated for a fish species, and are in keeping with the long and intensive breeding strategy observed in the wild population.     

Reproductive effort and associated costs in bison (Bison bison): do older mothers try harder?

To examine the prediction that reproductive effort should increasewith age, I compared female bison (Bison bison) of varying agesin terms of maternal behavior, reproductive costs, and the qualityand quantity of offspring. In support of predictions, oldermothers showed more tolerance toward and spent more time nursingtheir calves in months 1–5 than expected based on agedifferences in size and presumed milk production. The two oldestmothers, who died before calving again, appeared to make thegreatest effort. When mothers were barren between calves, whichoccurred more often in older females, their increased investmentenhanced the weight, dominance, maturation rate, and/or fecundityof previous and subsequent offspring. In contrast to predictions,however, offspring quality did not generally increase with maternalage; in fact, primiparae tended to produce larger daughters.Nursing behavior varied little in month 1, when offspring conditionwas apparendy most affected. Young mothers contacted their calvesmore often and tended to graze more during the first month.Reproductive costs appeared greater for young mothers; theirfecundity in years after calving was relatively low. Proportionalweight loss following reproduction decreased with age, althoughchanging growth rates complicate interpretation. The resultssuggest a general decrease in reproductive effort, or at leastin associated costs, during the reproductive life span. Thispattern may be due in part to the increased experience and dominancestatus of older mothers, through which they may avoid some ofthe reproductive costs incurred by primiparae.     

Reproductive effort,fecundity and energy allocation in two species of the genus Perinereis (Polychaeta: Nereididae)

Summary

The reproductive effort in terms of fecundity and energy allocation was studied in two species of semelparous polychaetes belonging to the genus Perinereis, living in the same environment, with different reproductive modalities. There is a great individual variability both in terms of reproductive effort and fecundity. Fecundity varied from 4080 to 15000 oocytes in P. rullieri and from 7000 to 26000 in P. cultrifera; no linear relationship was found between oocyte number and total jaw length utilised as size index. The energy content of germinal and somatic tissues was determined by Differential Scanning Calorimeter (DSC). The reproductive effort was calculated as RE = E_G/(E_G + E_S) where E_G is the total energy in germinal tissues and E_S is the total energy in somatic tissues. Reproductive effort is very high with mean values of 0.62 for P. rullieri and 0.79 for P. cultrifera. The different amounts of energy allocated in germinal tissues can be attributed to the different reproductive modalities—P. rullieri reproduces in the atokous phase whereas P. cultrifera has conserved epitoky in its life-cycle. The lack of correlation between reproductive effort and size index strongly suggests that reproductive allocation does not increase with age. In semelparous species the variability in fecundity and reproductive effort observed cannot be interpreted in terms of a trade-off between fecundity and survival as in iteroparous species. In fact, in semelparous an individual allocates all available resources to reproduction and then dies.     

Reproductive effort of winkles (Littorina spp.) with contrasted methods of reproduction

Summary Reproductive effort was compared in Littorina rudis (ovoviviparous), L. nigrolineata (benthic eggs with direct development), L. neritoides and L. littorea (planktonic eggs and larvae). Three indices of reproductive effort were used: the proportion of total production committed to reproduction per unit time, the cumulative proportion of total production committed to reproduction up to a given age, and the ratio of reproductive production per unit time to somatic biomass at the beginning of the unit time interval. The indices were plotted against age and the snails ranked in order of their reproductive efforts at equivalent ages. When plotted on axes of absolute time, all three indices ranked the snails in the order L. neritoides. The rank order of reproductive efforts on a time axis standardized for growth rate was L. littorea, and on a time axis standardized for generation time was L. neritoidesL. rudis. The implications and relative merits of the different expressions of reproductive effort are discussed. No general relationship was found between the magnitude of reproductive effort and either reproductive type or population ecology among the Littorina spp., or among other intertidal prosobranchs reviewed from the literature. Semelparous species had among the highest reproductive efforts, but there was no clear separation in this respect from certain iteroparous species. Parental investment, measured as the energy content of an egg plus any accompanying jelly or capsular material, was one to two orders of magnitude greater in the species with direct development than in those with indirect development. There were significant differences between egg and hatchling sizes of L. rudis from three contrasted types of shore and these differences were thought to be correlated with the intensity of desiccation and predation on hatchlings rather than with demographic factors.     

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.     

Reproductive effort and costs of reproduction do not explain female-biased sex ratios in the moss Pseudocalliergon trifarium (Amblystegiaceae)

A fundamental assumption in life-history theory is that reproduction is costly. Higher reproductive investment for fruits than for flowers may result in larger costs of reproduction in females than in males, which is often used to explain male-skewed sex ratios in unisexual seed plants. In contrast, bryophytes have predominantly female-biased sex ratios, suggested to be a product of a higher average cost of sexual reproduction in males. Empirical evidence to support this notion is largely lacking. We investigated sex-specific reproductive effort and costs in the unisexual moss Pseudocalliergon trifarium that has a female-dominated expressed sex ratio and rarely produces sporophytes. Annual vegetative segment mass did not differ among male, female, and non-expressing individuals, indicating that there was no threshold-size for sex expression. Mean and annual mass of sexual branches were higher in females than in males, but branch number per segment did not differ between sexes. Prefertilization reproductive effort for females was significantly greater (11.2%) than for males (8.6%). No cost for sexual branch production in terms of reduced relative vegetative growth or decreased investment in reproductive structures in consecutive years was detected. A higher realized reproductive cost in males cannot explain the unbalanced sex ratio in the study species.     

Biogeography in the Death Valley region: evidence from springsnails (Hydrobiidae: Tryonia)

Allozyme and mitochondrial DNA variation were analysed to examine evolution of the nine species of springsnails (genus Tryonia) living in the Death Valley system (Owens and Amargosa basins) of southeastern California and southwestern Nevada. Both allozyme and mtDNA evidence indicate that this highly endemic fauna is non-monophyletic. Species from the upper Amargosa basin comprise a clade most closely related to snails living in the Colorado basin. Snails from the lower Amargosa basin (Death Valley trough) reflect a complex evolutionary history and two of these species are more closely related to an estuarine species from western California than to other snails of the region. These results indicate a commonality of pattern with the well-studied Death Valley pupfishes (Cyprinodon), which also are non-monophyletic and include species that are most closely related to Colorado basin congeners. These biogeographic patterns are interpreted within the context of a recently proposed model for the early history of the lower Colorado River.