Egg trading in the simultaneously hermaphroditic polychaete worm Ophryotrocha gracilis (Huth)

Ophryotrocha gracilis (Huth 1934) is a simultaneously hermaphroditicpolychaete worm with external fertilization and a brief protandrousphase. The mating system of this species seems to meet conditionsleading to the establishment of egg-trading behavior. Experimentsshowed that mating occurs in pairs composed of two simultaneoushermaphrodites; sex roles are sequentially alternated and self-fertilizationis avoided. Egg reciprocation is kept evolutionarily stableby laying eggs in multiple, small-sized egg clutches and reducingthe reproductive success of pairs in which one of the two partnersdoes not reciprocate egg exchange. The frequency of ovigeroushermaphrodites in mass cultures of O. gracilis is about 50".Such a high mate availability preadapts hermaphrodites of O.gracilis to change partners very frequently and to reduce investmentin parental care, contrary to what is observed in another egg-trading,simultaneously hermaphroditic species, O. diadema. Laboratorypopulations of O. diadema have a frequency of only 17% ovigeroushermaphrodites     

Partner fidelity and egg reciprocation in the simultaneously hermaphroditic polychaete worm Ophryotrocha diadema

The mating system of the simultaneously hermaphroditic polychaeteworm Ophryotrocha diadema consists of a regular egg exchangebetween partners of a pair. Such reciprocal egg exchange hasbeen considered a form of cooperation, where one partner cooperatesby offering its eggs to be fertilized and expects to receivepartner's eggs to fertilize. Frequency of cases in which hermaphroditescheated (i.e., failed to give up eggs at their turn) and responsesto cheating were estimated by analyzing the behavior of 38 triplets of ovigerous hermaphrodites over a 2-week period. Thepartner did not reciprocate 8% of egg layings. The cheatedpartner did not detect most cases of cheating (16 out of 25).Such a low frequency of cheating can explain why no retaliationmechanism evolved in this species. Sixty-eight percent of the individuals from the original pairs deserted even if their partnersnever cheated them; therefore, cheating cannot be consideredthe cause of desertion. Rather, desertion appeared to be aconsequence of availability of a new partner whose oocyteswere riper than those of the old partner. It occurred becausethe opportunity arose for an immediate reward, indicating thatO. diadema egg exchange differs from that originally describedin some serranid fish as egg trading. The relationship betweencosts of desertion and population size is discussed.     

Life history and sex allocation in the simultaneously hermaphroditic polychaete worm Ophryotrocha diadema: the role of sperm competition

Sex allocation theory predicts that, in hermaphroditic organisms,individuals allocate a fixed amount of resources divided amongmale and female functions to reproduction and that the proportiondevoted to each sex depends on the mating group size. As themating group size increases, hermaphrodites are predicted toallocate proportionally more resources to the male and lessresources to the female function (approaching equal allocationto both sexes) to face increased sperm competition. Up to nowlittle experimental evidence has been provided to support thetheory in hermaphroditic animals. Facultative shift betweenmale and female allocation in response to variation in localgroup size does occur in several taxa but not always in theexpected direction and not with similar patterns. In the protandricand then simultaneously hermaphroditic polychaete worm Ophryotrochadiadema reproductive resources are flexibly allocated in theprotandrous and the hermaphroditic phase. The cost of male reproductionduring adolescence is spread over the whole energy budget ofthe animal as shown by the shortening of lifespan and the loweringof growth rate in individuals with enhanced male expenditureduring the protandrous phase. Moreover, in this species, shortterm sex allocation adjustments differ from those describedin other taxa. Individuals regulate their reproductive outputso that where reproductive competitors are present, the numberof female gametes is strongly reduced but the number of malegametes (although it changes) is not significantly increased.Resources subtracted from the female function are not directlyallocated to sperm production, but to expensive male behaviorsthat are likely to enhance male reproductive success. Theseresults are discussed in the light of the relevance of sexualselection in large populations of hermaphrodites.     

Life history trade-offs and response to selection on egg size in the polychaete worm Hydroides elegans

In order to examine the genetic relationships among life-history traits in a hermaphroditic species we used artificial selection for increased egg size and measured correlated responses across the life cycle of the serpulid polychaete Hydroides elegans, a protandrous sequential hermaphrodite. We recorded sex ratios across generations, and measured egg size, egg energy, larval volume at two time points, juvenile tube length, adult dry weight and fecundity after selection. Selection for larger eggs produced positive correlated responses in egg energy, fecundity and larval size at competence. Selection for increased egg size was also manifested by earlier sex change and this resulted in selected individuals spending less time as males relative to controls. We propose that egg size is negatively correlated with duration of andromorphy, that is, that female fitness trades off with male fitness.     

Heritable variation of sex ratio in a polychaete worm

Ophryotrocha labronica is a gonochoristic polychaete worm whose sex determining mechanism and sex ratio control are supposed to be polygenic. From a lab population, whose sex ratio (i.e., proportion of males) was 0.5, the estimate of sex ratio heritability by offspring-father regression was 0.54 ± 0.15 and by offspring-mother regression was not significantly different from 0. Estimate of sex ratio repeatability between successive broods of a pair was 0.64 ± 0.33. Since female parents do not contribute in any way to the variability of sex ratio, sex ratio variation seems to be largely a paternal character. On the basis of these estimates we advance the hypothesis that in this species sex is determined by a multilocus genetic system, allowing the combined effects of a female major sex gene (which could give rise to a form of female heterogamety) and masculinizing modifiers. The hypothesis that the male sex has the least canalised sexual differentiation is supported by the observation that some old males developed oocytes.     

Sperm trading in a hermaphroditic flatworm: reluctant fathers and sexy mothers

When matings are frequent and received sperm are digested, hermaphrodites should trade sperm when mating. We investigated sperm trading in the flatworm Schmidtea (Dugesia) polychroa and manipulated mating interests to investigate its possible causes. In 106 mating pairs consisting of nonisolated individuals, no sperm donation in either direction (35%) and reciprocal exchange (38%) were more common than expected by chance, whereas unilateral transfer (27%) was less frequent, confirming sperm trading. The amount of sperm donated depended on the availability of self-sperm, not on the amount received. Animals with more allosperm from previous matings had more self-sperm and consequently donated more. This suggests that sperm digestion boosts sperm production. In a second experiment, 'mixed-interest' pairs consisting of a nonisolated (N) and an isolated individual (I), NxI, were compared with IxI and NxN pairs. Whereas IxI pairs were eager and NxN reluctant to mate, NxI pairs showed an intermediate likelihood of mating. Whereas NxN pairs traded sperm, the other two groups did not. The change in behaviour in N individuals in the NxI treatment suggests precopulatory assessment and mating in relation to phenotypic mate quality. Isolated individuals are attractive, presumably because they donate large sperm clumps unconditionally and contain fewer allosperm, implying reduced sperm competition. The reduced reluctance in N individuals to mate with, and to inseminate, previously isolated partners suggests that female quality is an important factor in male sperm donation decisions. Hence, S. polychroa may be choosier than previously assumed. Copyright 2000 The Association for the Study of Animal Behaviour.     

Novel organelle associations in photoreceptors of a serpulid polychaete worm

Cerebral and branchial eyes of serpulid polychaetes have been studied by electron microscopy for the first time. In one species both eye types possess a novel ultrastructure. The receptor cells of the simple cerebral ocelli are rhabdomeric and display a close structural and functional relationship between ciliary rootlets and mitochondria. The receptors of the compound branchial eyes contain both a rhabdom and a stack of photosensitive ciliary membranes.     

Ecdysone receptor homologs from mollusks, leeches and a polychaete worm

The genomes of the mollusk Lottia gigantea, the leech Helobdella robusta and the polychaete worm Capitella teleta each have a gene encoding an ecdysone receptor homolog. Publicly available genomic and EST sequences also contain evidence for ecdysone receptors in the seahare Aplysia californica, the bobtail squid Euprymna scolopes and the medicinal leech Hirudo medicinalis. Three-dimensional models of the ligand binding domains of these predicted ecdysone receptor homologs suggest that each of them could potentially bind an ecdysone-related steroid. Thus, ecdysone receptors are not limited to arthropods and nematodes.     

Conspecific brood parasitism and egg rejection in Great-tailed Grackles

In this study, we tested whether conspecific brood parasitism (CBP) has selected for egg rejection behavior in the colonial Great-tailed Grackle Quiscalus mexicanus . No evidence of CBP was recorded at 797 Great-tailed Grackle nests, and we did not induce CBP by experimentally removing nests while grackles were laying. We determined experimentally that Great-tailed Grackles are determinate layers, an attribute opposite to that sometimes associated with CBP. Despite the absence of CBP, Great-tailed Grackles rejected 8% of experimentally introduced conspecific eggs, rarely rejecting or damaging their own eggs. Conspecific eggs added to nests during incubation tended to be rejected more frequently than eggs switched between nests, and eggs that differed the most from the host's eggs tended to be rejected sooner. There was no relationship between rejection and the stage of the nest cycle when experimental parasitism occurred; however, eggs were rejected faster when added during the prelaying and incubation stages than during laying. Evidence suggests, therefore, that egg rejection behavior in Great-tailed Grackles has not evolved in response to CBP.     

Characterization of novel F-actin envelopes surrounding nuclei during cleavage of a polychaete worm

F-actin accumulations and their possible functions were investigated during cleavage of the polychaete Ophryotrocha puerilis. Unusual cytoplasmic accumulations of F-actin were detected which have never been described before in animal embryos. As shown by TRITC-phalloidin labeling, envelopes of F-actin surrounded late prophase nuclei for a short period of time. DTAF-immunofluorescence of beta-tubulin showed that the F-actin envelope was closely associated with microtubules of the developing spindle apparatus. However, experimental disassembly of microtubules by nocodazole did not prevent the assembly of the F-actin envelope. Disturbance of F-actin envelope formation by cytochalasin B did not alter the course of mitotic events, i.e. position of the nuclei and orientation of the spindle apparatus were not affected, although the respective blastomeres remained uncleaved. However, disassembly of the F-actin envelope correlated temporally with breakdown of the nuclear envelope. Therefore, it is suggested that this new structure plays a role in fragmentation of the nuclear envelope during cleavage of Ophryotrocha puerilis.     

Sensory innervation of the somite in the polychaete worm Nephthys hombergii]

Studies have been made on the peripheral sensory innervation of N. hombergii stained with methylene blue. It was shown that sensory elements of the polychaete are presented by several types of nervous cells which are symmetrically embedded between the circular and longitudinal muscle layers in each of the segments. Peripheral projections of these cells terminate in the epithelium, on muscle fibers and in the connective tissue. Central projections pass to the abdominal nerve chain. All. the observed sensory cells have a constant localization and shape, the latter presumably being determined by their polyvalent functions.     

Conspecific brood parasitism and egg quality in blue tits Cyanistes caeruleus

Laying eggs in nests of unrelated conspecific pairs to parasitize their parental care is a common phenomenon in birds. In blue tits Cyanistes caeruleus such conspecific brood parasitism (CBP) has never been reported in the literature. However, in a situation where breeding density was extremely high, we found six nests to be parasitized with eggs of conspecific females. Natural selection may favour elevated competitiveness of parasite young, since the negative consequences of increased sibling competition are incurred on the unrelated host parents and siblings, and therefore do not act as inclusive fitness costs for the parasites. Parasitizing females could achieve such a competitive advantage for their offspring by laying larger eggs or eggs with higher concentrations of testosterone in the yolk. We analyzed these parameters of the six parasitized nests, but did not find that parasite eggs differ systematically in these aspects from host eggs, nor that parasite eggs showed resemblance to host eggs. We suggest that a shortage of available nest sites caused some females to use CBP as a best-of-a bad job strategy, but that either the occurrence of CBP is too rare to lead to strong selection for egg adjustments or that parasitizing females are unable to do so.     

How to evade a coevolving brood parasite: egg discrimination versus egg variability as host defences

Arms races between avian brood parasites and their hosts often result in parasitic mimicry of host eggs, to evade rejection. Once egg mimicry has evolved, host defences could escalate in two ways: (i) hosts could improve their level of egg discrimination; and (ii) negative frequency-dependent selection could generate increased variation in egg appearance (polymorphism) among individuals. Proficiency in one defence might reduce selection on the other, while a combination of the two should enable successful rejection of parasitic eggs. We compared three highly variable host species of the Afrotropical cuckoo finch Anomalospiza imberbis, using egg rejection experiments and modelling of avian colour and pattern vision. We show that each differed in their level of polymorphism, in the visual cues they used to reject foreign eggs, and in their degree of discrimination. The most polymorphic host had the crudest discrimination, whereas the least polymorphic was most discriminating. The third species, not currently parasitized, was intermediate for both defences. A model simulating parasitic laying and host rejection behaviour based on the field experiments showed that the two host strategies result in approximately the same fitness advantage to hosts. Thus, neither strategy is superior, but rather they reflect alternative potential evolutionary trajectories.     

Cytochemical localization of acid phosphatase in light- and dark-adapted eyes of a polychaete worm,Nereis limnicola

Summary The amount and distribution of the lysosomal enzyme acid phosphatase in light- and dark-adapted eyes of the brackish-water annelid Nereis limnicola were studied by standard cytochemical techniques. Precipitate from the acid phosphatase reaction was observed in Golgi-endoplasmic reticulum-lysosomal complexes, primary lysosomes, and secondary lysosomes, formed by fusion of primary lysosomes with phagocytic and pinocytic vesicles containing products of presumed rhabdomeric degradation. The acid phosphatase reaction occurred in these organelles in both sensory and supportive cells of both light- and darkadapted ocelli. Secondary lysosomes were more abundant in sensory cells of illuminated ocelli than in those maintained in the dark. Sparse reaction product was found in Golgi cisternae, none in rough endoplasmic reticulum. We suggest that the increase of lysosomal activity in light-adapted eyes is correlated with the breakdown of photosensory microvilli upon exposure to light. A diagram of our interpretation of recycling of photoreceptoral membrane in N. limnicola is presented.     

The worm re-turns: hiding behavior of a tube-dwelling marine polychaete, Serpula vermicularis

The polychaete worm Serpula vermicularis (Serpulidae) filterfeeds at the mouth of its calcareous tube, but retreats intothe tube when startled by mechanical stimuli likely to be associatedwith predators. While in its tube, a worm is safe but cannotfeed. Thus, hiding has a lost-opportunity cost. We show thatthis cost can be substantial, given that food in the naturalhabitat appears in pulses, and good feeding conditions may notlast long or recur frequently. We expect that a worm's hidingtime will be sensitive to the lost-opportunity cost, and wepresent data from a series of experiments that support thisprediction. The worms seem able to track relatively short-termchanges in food availability, and some evidence suggests thatthey assess food availability on a relative basis, comparingcurrent feeding conditions to those recently experienced. Hidingand other types of cryptic behavior are common antipredatortactics, and animals may commonly adjust the durations of suchbehaviors to current benefits and costs (including lost opportunity),as they perceive them     

The evolution of egg size in the brood parasitic cuckoos

We compared genera of nonparasitic cuckoos and two groups ofparasitic cuckoos: those raised together with host young ("nonejectors")and those in which the newly hatched cuckoo either ejects thehost eggs or chicks, or kills the host young ("ejectors"). Nonejectorsare similar to their hosts in body size and parasitize largerhosts than do ejectors, which parasitize hosts much smallerthan themselves. In both types of parasite, the cuckoo's eggtends to match the host eggs in size. To achieve this, nonejectorshave evolved a smaller egg for their body size than have nonparasiticcuckoos, and ejectors have evolved an even smaller egg. Amongejector cuckoo genera, larger cuckoos have larger eggs relativeto the eggs of their hosts, and the relationship between cuckooegg volume (mass of the newly-hatched cuckoo) and host egg volume(mass to be ejected) did not differ from that predicted by weight-liftingallometry. However, comparing among Cuculus cuckoo species,the allometric slope differed from the predicted, so it is notclear that egg size is related to the need to give the cuckoochick sufficient strength for ejection. Comparing the two mostspeciose ejector genera, Chrysococcyx cuckoos (smaller and parasitizedome-nesting hosts) lay eggs more similar in size to their host'seggs than do Cuculus cuckoos (larger and parasitize open cup–nestinghosts). Closer size-matching of host eggs in Chrysococcyx mayreflect the following: (1) selection to reduce adult body massto facilitate entry through small domed nest holes to lay, and(2) less need for a large egg, because longer incubation periodsin dome-nesting hosts allow the young cuckoo more time to growbefore it need eject host eggs.     

Monophyletic origin of brood care in damselfishes

The absence of a pelagic larval stage and brood care has evolved very few times in coral reef fishes. Damselfishes, a widely represented group with more than 380 species, includes only three such species, the monotypic Acanthochromis polyacanthus, and the two Altrichthys species, Altrichthys azurelineatus and Altrichthys curatus. In a recent study, Cooper et al. provided a comprehensive phylogenetic hypothesis for the damselfish family, based on mitochondrial and nuclear sequences, with more than 100 species that included all extant genera (Cooper et al., 2009). A. polyacanthus and sequences from formalin-preserved tissue of A. curatus did not cluster together, indicating that brood care may have evolved independently twice in damselfishes. Here, we use mitochondrial and nuclear molecular markers from fresh specimens of both Altrichthys species. We found that Altrichthys and Acanthochromis are closest relatives, which suggests that brood care may have evolved only once in damselfishes. Due to the limited geographic range of Altrichthys (western Philippines) and the abundant presence of Acanthochromis in that region, we also suggest that brood care may have originated in that region.     

The timing of spawning and egg production as constraints on male mating success in a simultaneously hermaphroditic fish

Synopsis The evolutionary stability of simultaneous hermaphroditism depends in part on the existence of constraints on the potential for male mating success. In the seabasses (Serranidae), several species of simultaneous hermaphrodites divide each day's clutch of eggs into parcels that are spawned sequentially and alternately with a partner. This behavior is thought to be one source of constraint on male mating success. A possible related source is the pattern of egg production. A study was therefore performed on the chalk bass,Serranus tortugarum, to examine this pattern. The results indicate that eggs are readied for spawning gradually over the course of the daily spawning period. The pattern of egg production acts jointly with spawning behavior in constraining male mating success. This pattern may be a pre-existing state to which the mating system has been adjusted, or it may have co-evolved with the mating system.