Evolution of egg size and fertilisation efficiency in sea stars: large eggs are not fertilised more readily than small eggs in Patiriella (Echinodermata: Asteroidea)

Fertilisation kinetics theory suggests that, when sperm are limiting, the larger eggs of broadcast-spawning marine organisms ought to be fertilised more frequently than smaller eggs, because they provide a bigger target for searching sperm. Whilst this effect has been demonstrated within species, it is not known if this pattern holds among species. We tested whether a large difference in egg size between congeneric seastars with contrasting planktotrophic and lecithotrophic modes of development results in differences in the likelihood of eggs being fertilised in sperm-limiting situations. Measurement of egg sizes and sperm swimming speeds led to the prediction that the sperm–egg collision rate constant for Patiriella calcar (420-µm-diameter egg) should be nine times greater than for P. regularis (140-µm-diameter egg). Although the eggs of P. calcar should be fertilised at greater rates in low sperm concentrations, they were not. When gametes were allowed to mix for 10 s, the hypothesis that P. calcar eggs required less sperm than P. regularis to ensure 50% of eggs were fertilised was rejected. When gametes were mixed for 5 min, P. regularis eggs were more frequently fertilised, but the difference was not statistically significant. We conclude there must be a difference between these species in the likelihood that when a sperm finds a conspecific egg it can successfully fertilise. This apparent uncoupling of egg size and likelihood of fertilisation suggests that fertilisation is not a major constraint on the evolution of egg size in these seastars.

Reproductive development, GnRHa-induced spawning and egg quality of wild meagre (Argyrosomus regius) acclimatised to captivity

The objective of the study was to acclimatise wild-caught meagre (Argyrosomus regius) to captivity to produce viable eggs for aquaculture production. Twelve meagre (3 males and 9 females, mean weight = 20 ± 7 kg) were caught and transported to a land-based facility on 26 October 2006. During, March to June 2007, all three males were spermiating and five of the nine females were in vitellogenesis with mean maximum oocyte diameter ≥550 μm. No spontaneous spawning was observed. Two hormone treatments, either a single injection of gonadotropin-releasing hormone agonist (GnRHa, 20 μg kg−1 for females and 10 μg kg−1 for males) or a slow-release implant loaded with the same GnRHa (50 μg kg−1 for females and 25 μg kg−1 for males), were used to induce spawning on three different dates on 26 March 2007, 4 May 2007 and 18 April 2008. From each spawning event, the following parameters were determined: fecundity, number of floating eggs, egg size, fertilisation and hatching success, unfed larval survival, and proximal composition and fatty acid profile of the eggs. In 2007, two females that were injected on 26 March and 4 May spawned a total of 5 times producing 9,019,300 floating eggs and a relative fecundity of 198,200 eggs kg−1 and two different females that were implanted on the same dates spawned 14 times producing 12,430,000 floating eggs and a relative fecundity of 276,200 eggs kg−1. In 2008, a pair that was implanted spawned five times producing a total of 10,211,900 floating eggs and a relative fecundity of 527,380 eggs kg−1. The latency period was 48–72 h. Parameters were compared between hormone treatments, date of hormone induction and parents determined by microsatellites. Percentage hatch and egg size were 70 ± 0.3% and 0.99 ± 0.02 mm, respectively, for GnRHa-implanted fish and were significantly higher (P < 0.05) compared to 30 ± 0.3% and 0.95 ± 0.03 mm, respectively, for injected fish. Few differences were observed in proximal composition and fatty acid profile and for all spawns mean (% dry weight) lipid content was 17.3 ± 3.0%, carbohydrate was 4.4 ± 1.9% and protein was 31.5 ± 6.4% and the essential fatty acids: Arachidonic acid (ARA, 20:4n-6) ranged between 0.9 and 1% (of total fatty acids), eicosapentaenoic acid (EPA 20:5n-3) 7.7–10.4% and docosahexaenoic acid (DHA 22:6n-3), 28.6–35.4%. All good quality spawns were obtained in the second and/or third spawn after GnRHa treatment, whereas all bad quality spawns were obtained either on the first spawn or after the fifth spawn. Both spawning protocols gave commercially viable (1,000,000+) numbers of good quality eggs that could form the basis of a hatchery production.

Sperm environment affects offspring quality in broadcast spawning marine invertebrates

The provisioning of offspring can have far-reaching consequences for later life in a wide range of organisms and generally this provisioning is thought to be under maternal influence or control. In experiments with a broadcast-spawning ascidian, we found that the size of offspring was determined by egg size and the abundance of sperm present during fertilization. Larger eggs were fertilized at low sperm concentrations, whilst smaller eggs were successfully fertilized at high sperm concentrations. These differences in fertilized egg size resulted in differences in the development rate, hatching success and mean size of the subsequent larvae. Our results suggest that, in contrast to females that reproduce by other mating systems, free-spawning mothers lack some control over the provisioning of offspring. Furthermore, because males can alter the sperm environment, they can exert paternal (non-genetic) control over key offspring characteristics.

Ecology of epifaunal caridean shrimps in the Hopkins River estuary, and the role of estuaries in the life history of the atyid "Paratya Australiensis" Kemp, 1917 in south-eastern Australia

This study examined the factors affecting the distribution and abundance of epifaunal caridean shrimps in seagrass meadows of the Hopkins River estuary in south-western Victoria, Australia, and investigated the life history patterns of the freshwater Parana australiensis, found for the first time in estuaries. Adult and sub-adult shrimps were surveyed in seagrass meadows along the estuary over two years, and their planktonic larvae were surveyed in adjacent waters. Three species were collected. The marine Palaemon serenus occurred only near the mouth, summer to autumn, in high salinities. The marine/estuarine Macrobrachium intermedium occurred throughout the estuary. Adults were most abundant in late autumn, and least abundant in summer (unlike trends reported in marine meadows). Densities were higher and less variable in downstream meadows. P. australiensis occurred in the upper estuary all year, most abundantly in spring, due to migration from the river after peak discharge. Ovigerous females dominated, while males, showing less migration into the estuary, dominated above estuarine influence. Adults disappeared from the estuary in summer as salinity rose. Breeding period for P. australiensis was briefer in the estuary (September-December) than upstream (July-April). M. intermedium began breeding later in the upper estuary (November/December-March) than in the lower estuary (October-March), probably reflecting a physiological response to lower salinity, rather than an interaction with P. australiensis. No ovigerous P. serenus were found in the estuary. Larvae of P. australiensis and M intermedium occurred abundantly throughout the estuary, but P. serenus larvae did not. P. australiensis was an early coloniser to the plankton after peak discharge (November-December). Larvae concentrated in the deep saline layer at the head of the intruding salt wedge, thus probably maintaining longitudinal position. Diurnal vertical migrations were evident within the salt wedge, and in a deep pool above tidal influence. M. intermedium larvae occurred October-May in the lower estuary and November-April in the upper estuary, peaking in abundance one to two months after P. australiensis. They were associated with low surface flows and surface salinities greater than 10, over an anoxic deeper layer. All three species exhibited extended development of euryhaline larvae in the laboratory. Tolerances and optimal salinities of larvae of the three species reflected their distributions. M. intermedium was the most euryhaline species. P. australiensis larvae were tolerant of higher salinities than juveniles of adults: capable of developing in salinity of at least 15. Most P. australiensis juveniles recruited to the estuary November-December, after which numbers declined dramatically. After settlement, most recruits probably migrated upstream out of the estuary. Two cohorts of M. intermedium recruited to the estuary from larvae in summer (December and February), but some juveniles also migrated from adjacent coastal waters. Post-larval migration was at least as important a determinant of abundance as direct recruitment from estuarine, planktonic larvae in all three species. Distributions among seagrass meadows along the estuary were determined primarily by physico-chemical patterns driven by hydrological changes. Seasonal variations in salinity and temperature were strongly associated with seasonal variations in shrimp abundance. Salinity tolerances of adults of the three species reflected their distribution patterns. Biotic interactions were more important in determining distributions within meadows. P. australiensis, when abundant, were associated with seagrass biomass. M. intermedium were also, but when seagrass was sparsest and least extensive. The two species apparently partitioned the seagrass meadow according to depth in early summer. Laboratory experiments suggested P. australiensis was displaced from deeper water by M. intermedium. Preference for vegetative complexity and competition for position within meadows suggest the underlying importance of predation in regulating shrimp populations. A survey of south-eastern Australian estuaries found P. australiensis larvae abundant in all stable, open, well-developed, salt-wedge estuaries where adults were abundant. Adults were most abundant in low salinities among submerged leafy macrophytes. Reproductive traits of P. australiensis were compared in estuarine and fresh reaches of three rivers. Early in the breeding season, egg size was smaller, and (size-specific) egg number larger in estuaries than upstream. A trade-off between egg size and egg number resulted in no difference in total (size-specific) reproductive investment between locations. Reproductive investment tended to decrease at some locations over the breeding season, and this decrease was a result of decreased egg size in most cases. The decrease in reproductive investment probably reflected reduced food availability for the adult, while the reduced egg size was probably a response to improved conditions for larval development. In the Hopkins River, larger egg size at upstream sites was reflected in larger early stage larvae. Later stage larvae were larger in the estuary, suggesting more favourable conditions for larval development. Allozyme electrophoresis showed the P. australiensis populations in each of the three rivers to be distinct. Allozyme frequencies were not different within the Hopkins River, but upstream and estuarine locations in the Curdies and Gellibrand were different. Although some variation in reproductive traits within catchments may have been due to genotypic differences, trade-offs between egg size and number, and decreases in egg size over summer were probably due to plastic responses to environmental cues. It is proposed P. australiensis inhabits and reproduces in both estuarine and freshwater environments by plastic response to environmental conditions. Recruitment to estuaries is dependent on the presence of suitable adult, littoral habitat, and a stable salt wedge for larval retention. Estuaries are important recruitment sites for P. australiensis, potentially allowing an extra brood each year before riverine recruitment. Estuarine broods could constitute a large part of the total fecundity of P. australiensis females. Euryhaline larvae and estuarine recruitment of P. australiensis suggest marine transport of larvae between estuaries as a possible dispersal mechanism for Paratya species.

Maternal effects in the Zebra Finch : a model mother reviewed

Birds are a particularly good group with which to examine the importance of maternal effects, as parental contributions can be relatively easily quantified compared with other groups. There have undoubtedly been more studies on maternal effects in the Zebra Finch than any other single bird species. Studies of this species have examined the importance of maternal effects mediated through sex allocation, size, nutrients and hormones of of eggs, incubation behaviour and provisioning levels. A synthesis of all of this work illustrates some contrasting results (e.g. many high-profile results have failed to be replicated), some very common patterns (e.g. investment shifts through the laying sequence), and potentially interesting and complex interactions between traits (e.g. between sex of offspring and hormonal profiles of eggs). This extensive collection of work on the Zebra Finch provides useful general insight into the patterns of maternal investment in birds and the effects on offspring phenotype. However, we caution that the literature is probably littered with studies that have overemphasised the importance of some maternal effects and recent studies have highlighted analytical and logical flaws that have probably led to misplaced confidence in some of the findings reported to date. Finally, it is worth considering that the bulk of the literature is based on studies of captive domesticated birds and ecological and physiological data from individuals in the wild is currently lacking. The biological relevance of maternal effects documented in this model species is therefore unclear.

Transfer of maternal antibodies against avian influenza virus in mallards (Anas platyrhynchos)

Maternal antibodies protect chicks from infection with pathogens early in life and may impact pathogen dynamics due to the alteration of the proportion of susceptible individuals in a population. We investigated the transfer of maternal antibodies against avian influenza virus (AIV) in a key AIV host species, the mallard (Anas platyrhynchos). Combining observations in both the field and in mallards kept in captivity, we connected maternal AIV antibody concentrations in eggs to (i) female body condition, (ii) female AIV antibody concentration, (iii) egg laying order, (iv) egg size and (v) embryo sex. We applied maternity analysis to the eggs collected in the field to account for intraspecific nest parasitism, which is reportedly high in Anseriformes, detecting parasitic eggs in one out of eight clutches. AIV antibody prevalence in free-living and captive females was respectively 48% and 56%, with 43% and 24% of the eggs receiving these antibodies maternally. In both field and captive study, maternal AIV antibody concentrations in egg yolk correlated positively with circulating AIV antibody concentrations in females. In the captive study, yolk AIV antibody concentrations correlated positively with egg laying order. Female body mass and egg size from the field and captive study, and embryos sex from the field study were not associated with maternal AIV antibody concentrations in eggs. Our study indicates that maternal AIV antibody transfer may potentially play an important role in shaping AIV infection dynamics in mallards.

Does female nuptial coloration reflect egg carotenoids and clutch quality in the Two-Spotted Goby (Gobiusculus flavescens, Gobiidae)?

Summary

1 - Carotenoid-based ornamentation has often been suggested to signal mate quality, and species with such ornaments have frequently been used in studies of sexual selection.

2 - Female Gobiusculus flavescens (Two-Spotted Goby) develop colourful orange bellies during the breeding season. Belly coloration varies among mature females, and previous work has shown that nest-holding males prefer females with more colourful bellies. Because males invest heavily in offspring during incubation, the evolution of this preference can be explained if colourful females provide males with eggs of higher quality.

3 - We tested this hypothesis by allowing males to spawn with ‘colourful’ and ‘drab’ females and comparing parameters including egg carotenoid concentration, clutch size, hatchability and larval viability between groups. We also investigated relationships between egg carotenoid concentration and clutch quality parameters.

4 - Eggs from colourful females had significantly higher concentrations of total carotenoids than drab females, and photographically quantified belly coloration was a good predictor of egg carotenoid concentration.

5 - Colourful females produced slightly larger clutches, but female belly coloration was not related to any measure of clutch quality. In addition, there were no significant relationships between egg carotenoids and clutch quality. Females with high levels of egg carotenoids spawned slightly earlier, however, possibly because they were more ready to spawn or because of male mate choice.

6 - Our results call into question the generality of a causal link between egg carotenoids and offspring quality.

Harem size and oviposition behaviour in a polygynous bark beetle

1. Harem polygyny can have fitness benefits and costs on females. In bark beetles of the genus Ips the latter may include within-harem competition between larvae. However, earlier competition between females for male care and mating opportunities may also influence oviposition behaviour. There has been relatively little investigation into the relationship between harem size and initial egg output. The present study investigated this relationship in the bark beetle Ips grandicollis.
2. The measure of egg output used was the number of eggs in the gallery with the most eggs in each harem. Mean ( ± SE) harem size of 242 observed harems was 3.25 ± 0.10. A curvilinear relationship was found between egg output and harem size, with females in smaller harems (one to four females) laying more eggs with increased harem size. However, females in larger harems (five to seven females) laid fewer eggs as harem size increased. The optimal harem size (in terms of number of eggs laid) was close to four females.
3. We found no evidence from a behavioural assay that females could preferentially choose unmated males over mated males with harems of two females. Additionally, the distribution of harem sizes suggests that females distribute themselves among males randomly.
4. The results suggest that harem size has effects on female reproduction that extend beyond larval competition and influence patterns of oviposition. The mechanism that determines why egg laying is greatest at intermediate levels is unknown. There is no evidence that smaller harems belong to lower quality males, but females may adjust egglaying behaviour in large harems as a result of reduced male attendance or anticipated larval competition.

The implications of adult morphology for clutch size in the flatback turtle (Natator depressa)

When the mean adult length and mean clutch volume of marine turtles are examined, a clear pattern for larger species to lay larger clutches is evident, in accord with predictions that female size constrains the available space for carrying eggs. However, when compared with this general trend, the volume of clutches laid by flatback turtles (Natator depressa) are smaller than expected. The implication is that the unusually flat morphology of flatback turtles, provides an additional constraint on their egg carrying capacity.

Reproductive investment and optimum clutch size of Loggerhead Sea Turtles (Caretta caretta)

(1) Data for loggerhead turtles (Caretta caretta L.) nesting on the Greek island of Cephalonia were used to develop a model which predicted the optimum clutch size. (2) There was a positive linear relationship between the number of clutches into which eggs could be divided and the total time spent by a nesting turtle on the beach, and hence a negative relationship between the time invested on the beach per egg and clutch size. (3) A previous study indicated that energy expenditure for nesting turtles on land is very high, so there may be a selective pressure to maximize clutch size in order to minimize the energy expended per egg laid. As there appeared to be no counterselective pressures favouring small clutches, clutch size should be constrained by a female's egg-carryingc apacity,w hich in turn could be expected to be related to her body size. Hence, a positive relationship between clutch size and body size was predicted, and was found in the population under study.

Resource allocation across the egg laying sequence in the wild zebra finch Taeniopygia guttata

Zebra finches have played a central role in the discovery of a variety of maternal effects over the past decade, with females shown to adjust resource allocation to their eggs in response to variables such as the appearance of their partner, their own condition, and the diet on which they are maintained. In addition to being the focus of some of the most high profile individual studies that have influenced maternal effects research in birds, the multitude of zebra finch studies together provide the most comprehensive set of data to illuminate general patterns and compare different maternally derived variables. Surprisingly, to date, virtually all of this work has focused on captive populations of the zebra finch that have been domesticated for many generations, and which are typically held under relatively constant environmental and dietary conditions. Here we report the first data on resource allocation across the egg laying sequence in a free-living wild population. Reassuringly we find that the patterns that have been found in the majority of studies of domesticated populations with respect to investment across the laying sequence were all present in the wild population. The size and mass of eggs increased through the laying sequence whilst the concentration of carotenoids significantly decreased across the laying sequence. Although there was no significant pattern with respect to testosterone across the laying sequence the first two eggs had a higher level of testosterone than the last few eggs in the clutch, which is also consistent with the findings of earlier studies in captive populations. © 2011 The Authors.