Lecithotrophic development and metamorphosis in the Indo-West Pacific brittle star Ophiomastix venosa (Echinodermata: Ophiuroidea)

Summary

The larval development of the ophiocomid ophiuroid Ophiomastix venosais described using SEM. The gastrula transforms into a uniformly ciliated early larva which progressively changes into a lecithotrophic late premetamorphic larva with a continuous bilateral ciliated band. This stage is short-lived and equivalent to a highly reduced ophiopluteus. Comparisons between O. venosa and other ophiuroid species whose development has been investigated suggest that, whatever the developmental mode (lecithotrophic or planktotrophic), a pluteus stage always occurs in ophiuroids with planktonic development. Two metamorphic stages were identified, the late metamorphic larva differing from the early one by the closure of the larval mouth. The appearance of the permanent mouth marks the end of the metamorphosis. The postlarva still possesses remnants of larval features. The transformation of the reduced ophiopluteus into a barrel-shaped metamorphic larva with transverse ciliated bands, a vitellaria larva, is followed. The possible occurrence of a unique type of metamorphic larva in non-brooding ophiuroids is discussed. Verification of this, however, needs further SEM investigations on metamorphic larva from species having “regular” planktotrophic development.     

Subsocial behaviour in Oniticellus cinctus (Coleoptera, Scarabaeidae): effect of the brood on parental care and oviposition

ABSTRACT. In Oniticellus cinctus (F.) the nest chambers each contain about twenty brood balls. Females enlarge the brood balls during the egg and larval stages and remain in the chamber for the whole period of brood development (1 month); they then make a new nest after 1 week. The presence of the brood releases parental care and ensures that the mother remains in the nest: she repairs defects in the brood balls and the nest, and expels other O. cinctus females. A new ball is formed around a naked O. cinctus larva, but larvae of other species are killed. In addition, the brood inhibits oviposition: removal (or addition) of brood balls stimulates (or inhibits) egg laying. In inhibited ovarioles, existing follicles are resorbed and production of new ones ceases. Control of clutch size by the brood is an adaptation to the nest structure and life history of O. cinctus. It may have an important role in the reproductive strategy of other insects with parental care.     

Comparative larval energetics of an ophiuroid and an echinoid echinoderm

The morphologically convergent larvae of the echinoderm classes Ophiuroidea and Echinoidea have been suggested to be functionally dissimilar when it comes to their capacities to feed, but little is known about whether these larvae are similar in terms of energetics. Here, we compare the energetics of early development of a tropical ophiuroid, Ophiocoma alexandri, and a temperate to tropical echinoid, Arbacia punctulata, two species with similarly sized eggs. Measurements of respiration and constituent analyses were performed on eggs and unfed larvae of both species. Members of both species showed an increase in oxygen consumption during morphogenesis followed by a lower, static rate once morphogenesis was complete (3 d for O. alexandri and 1.3 d for A. punctulata). Compared to the echinoid larvae, the ophiuroid larvae developed more slowly and had peak respiration rates that were 3.1× lower. Eggs of O. alexandri contained significantly more protein and significantly less triacylglycerol than eggs of A. punctulata. Energy utilization, as calculated via respiration measurements, closely matched decreases in energy content from the eggs to larvae as measured with biochemical constituent assays. Larvae of A. punctulata used 1.4× more energy to reach the pluteus stage than larvae of O. alexandri, and used 4× more energy during the first 9 d of larval life. These data suggest that echinoid larvae require more energy to develop to the feeding stage than ophiuroid larvae, and likewise have higher requirements for maintenance metabolism. Ophiuroid larvae may be more tolerant of low food levels due to their very low metabolic rates, but this advantage may be offset by their slower rate of development.     

E-β-ocimene, a volatile brood pheromone involved in social regulation in the honey bee colony (Apis mellifera)

BACKGROUND: In honey bee colony, the brood is able to manipulate and chemically control the workers in order to sustain their own development. A brood ester pheromone produced primarily by old larvae (4 and 5 days old larvae) was first identified as acting as a contact pheromone with specific effects on nurses in the colony. More recently a new volatile brood pheromone has been identified: E-β-ocimene, which partially inhibits ovary development in workers. METHODOLOGY AND PRINCIPAL FINDING: Our analysis of E-β-ocimene production revealed that young brood (newly hatched to 3 days old) produce the highest quantity of E-β-ocimene relative to their body weight. By testing the potential action of this molecule as a non-specific larval signal, due to its high volatility in the colony, we demonstrated that in the presence of E-β-ocimene nest workers start to forage earlier in life, as seen in the presence of real brood. CONCLUSIONS/SIGNIFICANCE: In this way, young larvae are able to assign precedence to the task of foraging by workers in order to increase food stores for their own development. Thus, in the complexity of honey bee chemical communication, E-β-ocimene, a pheromone of young larvae, provides the brood with the means to express their nutritional needs to the workers.     

Adaptational Significance of the Patterns of Ophiuroid Development

There is only limited information on ophiuroid developmentalpatterns, larvae being known for only 4% of the 2000 livingophiuroids. Three modes of development are distinguished (planktotrophic,direct, abbreviated) on the basis of egg size and number, larvalform and life span, and post-larval size. This generalizationis applied to an analysis of the larval form(s) of deep-seaophiuroids and the ancestral ophiuroid larva. Quantitative informationon rate of development and reproductive period, which has beenlacking for viviparous ophiuroids, is presented for Axiognathus( = Amphipholis) squamatus on the basis of original size-frequencydata. Reproduction is seasonal in the population studied anddevelopment to metamorphosis takes no longer than for planktotrophicspecies though the brooding period ranges from 3 to 7 months.Additional data indicate post-larvae of planktotrophic specieshave fewer arm segments than newly-hatched viviparous young,supporting the hypothesis that large size of juveniles is characteristicand selectively advantageous for viviparous ophiuroids.     

Distribution of Japanese temperate bass, <Emphasis Type="Italic">Lateolabrax japonicus</Emphasis>, eggs and pelagic larvae in Ariake Bay

We collected eggs and larvae of the Japanese temperate bass, Lateolabrax japonicus, and present horizontal and temporal changes of distribution relative to development and growth during the species pelagic life history in Ariake Bay. Sampling was conducted from the inner to central region (11 sampling stations) of Ariake Bay using a plankton net (80 cm diameter, 0.5-mm mesh) from November 2000 to February 2001. Both eggs and larvae were collected most abundantly in mid-December. The CPUE of eggs in the surface layer was higher than the middle layer, which is in contrast to that at the larval stage. Most eggs were collected around the central and western regions of the bay. The distribution of eggs shifted vertically to the middle layer with development. Yolk-sac larvae were collected in the central region of the bay, and preflexion and flexion larvae were more abundantly collected in the inner region of the bay. The body length of larvae around the inner bay was larger than in the central region. The pelagic life history can be summarized as follows: eggs are distributed around the central region of the bay and eggs and larvae expand their distribution to the inner and shallower waters with growth. We conclude that the shift of vertical distribution in pelagic stages and the hydrographic features of the middle layer form one of the mechanisms enabling the inshore migration of L. japonicus.     

Resource rivalry between brood mates of a facultative gregarious parasitoid Dendrocerus carpenteri

Larvae of Dendrocerus carpenteri Curtis (Hymenoptera: Megaspilidae) develop as solitary ectoparasitoids on the prepupae and pupae of primary aphid parasitoids inside the aphid mummy. First instars are aggressive and kill potential competitors; however, facultative gregarious development of two, and occasionally three, larvae may occur under superparasitism. To test the hypothesis that gregarious larvae share host resources equally, adult dry mass is compared between three brood types: ‘double’ mummies containing (i) two males; (ii) two females; or (iii) one male plus one female, respectively. Resource rivalry varies with the sex of the competing larvae. Surviving adults differ significantly in size if both wasps are of the same sex, male or female. A minimum amount of resources is required for a larva to be viable; this threshold does not differ between sexes and is independent of the sex of a competing larva. The outcome of competition between a male larva and a female larva varies with the amount of the available resources, with neither sex being inherently dominant over the other. Females are eight times more likely to be larger than a male competitor if the amount of host resources is ≥0.116 mg, whereas males can win over a female competitor in terms of adult size if the available resources are <0.116 mg. It is suggested that rivalry between larvae for limiting host resources constrains the transition from solitary to gregarious development and should be considered in studies of parasitoid life‐history evolution.     

Development and metamorphic loss of the musculature in larvae of the nudibranch Phestilla sibogae: A functional ontogeny

Developmental programmes for many marine invertebrates include the assembly of muscular systems appropriate to the functions of swimming and feeding in pelagic larvae. Upon metamorphosis, that musculature is often radically re-organized to meet very different demands of post-larval life. To investigate the development and fate of musculature in the nudibranch Phestilla sibogae, embryos, larvae and metamorphosing stages were fixed, labelled with phalloidin and examined with confocal microscopy. The resultant images revealed the sequential development of both large retractor muscles and numerous finer muscles that allow the larva to manipulate the velum, foot and operculum. Observations of living specimens at the same stages as those fixed for microscopy revealed the actions of the muscles as they developed. During metamorphosis, muscles with shell attachments disintegrate as the larva transforms into a shell-less juvenile. Notably, the massive velar, pedal and opercular retractor muscles disappear during metamorphosis in a sequence that corresponds to their loss of function. Other muscles, however, that appear to be important to the embryo and free-swimming larva persist into juvenile life. The comprehensive and detailed observations of the musculature presented here provide a solid foundation for comparisons with other species with different phylogenies and life histories.     

Facultative Vivipary is a Life-History Trait in Caenorhabditis elegans

Organisms partition their resources among growth, maintenance, and reproduction and, when resources become limiting, the allocation to one process necessitates reduced allocation to others. When starved, Caenorhabditis elegans adults retain progeny internally which then consume the parent body contents, and some of those larvae use the resources to reach the resistant, long-lived dauer stage. If starved under similarly extreme conditions, larvae from eggs laid outside of the body are unable to develop into dauers. We interpret this switch from ovipary, or laying eggs, to bearing live young as facultative vivipary. This switch is induced by starvation of late fourth-stage larvae, young adults, or gravid adults. In C. elegans, vivipary is the altruistic allocation of all available parental energy and nutrients to progeny, with the associated costs to adult hermaphrodites of truncated life span and fecundity. As a life-history trait, facultative vivipary is a survival-enhancing response to stress that may provide insights into the evolution of reproduction and longevity.     

Reproductive and subsocial behaviour in the ovoviviparous leaf beetle Gonioctena sibirica (Coleoptera: Chrysomelidae)

Abstract.

• 1 In the spring, females of the leaf beetle Gonioctena sibirica deposited larvae on the ventral surface of growing young leaves situated on the apical position of shoots of the willow Salix bakko.
• 2 The parent females remained with the larvae usually on the underside of the basal part of leaves, facing toward the base of shoots. When other arthropods approached, the females temporarily moved towards these intruders, showing aggressive behaviour such as swinging the body or stamping the legs. Many females remained with their larvae until the larvae grew into the final (fourth) instar. No female produced an additional brood in the field.
• 3 Broods from which parent females were experimentally removed suffered higher mortality than those in which females were left intact. Arthropods such as spiders and ants were observed preying on the larvae. In contrast, the survivorship of broods from which females were removed and intruders were excluded with a sticky substance applied to the base of twigs was not different from that of control broods. These results demonstrate that the main mortality factor of offspring is pedestrian arthropod predators and females physically repel the predators.
• 4 Potentially alternative reproductive strategies, such as producing a large number of offspring by iteroparity and/or larger brood(s) with less or no care, seem to be inhibited in G.sibirica by larval dependence on growing young leaves which are temporally limited and by ovoviviparity which may have limited brood size.

     

Evidence for Cohesive Dispersal in the Sea

As with many marine species, the vast majority of coral-reef fishes have a bipartite life cycle consisting of a dispersive larval stage and a benthic adult stage. While the potentially far-reaching demographic and ecological consequences of marine dispersal are widely appreciated, little is known of the structure of the larval pool and of the dispersive process itself. Utilizing Palindrome Sequence Analysis of otolith micro-chemistry (PaSA;) we show that larvae of Neopomacentrus miryae (Pomacentridae) appear to remain in cohesive cohorts throughout their entire pelagic larval duration (PLD; ∼28 days). Genetically, we found cohort members to be maternally (mtDNA) unrelated. While physical forcing cannot be negated as contributing to initial cohort formation, the small scale of the observed spatial structure suggests that some behavioral modification may be involved from a very early age. This study contributes to our ongoing re-evaluation of the processes that structure marine populations and communities and the spatial scales at which they operate.     

Length of pelagic period in echinoderms with feeding larvae from the Northeast Pacific

Minimum and maximum times from fertilization to settling are reported for larvae reared in the laboratory from 12 species of echinoderms representing four classes. For the holothuroid Parastichopus californicus (Stimpson) the maximum time observed was 13 weeks; for the asteroids Luidia foliolata Grube, Pisaster ochraceus (Brandt), and Pycnopodia helianthoides (Brandt) the maxima were 20, 32, and 21 weeks, respectively; for the ophiuroid Ophiopholis aculeata (Linnaeus) the maximum was 31 weeks; for the echinoids Brisaster latifrons (A. Agassiz), Dendraster excentricus (Eschscholtz), Allocentrotus fragilis (Jackson), and Strongylocentrotus droebachiensis (O. F. Müller) maxima were 24, 23, 36, and 21 weeks, respectively. The maximum pelagic periods in each class of echinoderms were approximately double other estimates based on cultured larvae. The pelagic periods in these cultures suggest a capacity for extensive dispersal in nature.     

Adult honeybee's resistance against Paenibacillus larvae larvae, the causative agent of the American foulbrood.

American foulbrood is a widespread disease of honeybee larvae caused by the spore-forming bacterium Paenibacillus larvae subsp. larvae. Spores represent the infectious stage; when ingested by a larva they germinate in the midgut. The rod-shaped vegetative forms penetrate the larva's intestinal tissue and start multiplying rapidly, which finally kills the larva. Spores fed to adult honeybees, however, do not harm the bees. We investigated this phenomenon. Specifically, we studied the influence of the adult honeybee midgut on the vegetative growth and on the germination of spores of P. larvae larvae. We focused on two groups of adult workers that are likely to have large numbers of spores in their gastrointestinal tracts in infected colonies: middle-aged bees, which are known to remove or cannibalize dead larvae and clean brood cells, and winterbees, which do not have frequent chances to defecate. We found that midgut extract from winterbees and worker-aged bees of different colonies almost completely inhibited the growth of the vegetative stage of P. larvae larvae and suppressed the germination of spores. The inhibiting substance or substances from the adult midgut are very temperature stable: they still show about 60% of their growth-inhibiting capacity against this bacterium after 15 min at 125 degrees C. We established a method to test growth-inhibiting factors against P. larvae larvae in vitro.     

转Bt基因抗虫棉对亚洲玉米螟抗性的研究

对转Bt抗虫棉苏抗103与GK—22对玉米螟抗性的研究结果显示：(1)用苏抗103嫩叶、簧,花、蕾饲曩第三代玉米螟幼虫,最终幼虫校正死亡率分别达100％、95．83％、83．33％与100％。校正化蛹下降率分别达100％、92．31％、50．07％与100％,校正幼虫成蛾减退率分别达100％、90．92％、0％与100％。(2)人工饲料中加入5个不同浓度苏抗103嫩叶饲喂第二代幼虫,除0．4％浓度,幼虫校正死亡率与成蛹下降率均为100％,0．4％浓度的幼虫期延长20．68％,蛹重下降18．24％,蛹期延长19．79％。(3)人工饲料中加入5个浓度的苏抗103或GK22嫩叶分别饲喂第三代幼虫,苏抗103上幼虫校正死亡率较第二代有所下降,分别为3．04％、-16．68％、28．2％、33．33和75％。GK22上幼虫校正死亡率分别为3．13％、26．89％、45．69％、70．816％和68．09％。其幼虫校正死亡率及幼虫期、蛹期、成虫校正减退率均有随浓度增加而上升的趋势。较亲本棚,对蛹重、雌成虫寿命、单雌产卵量、卵孵化率也有一定程度的影响。上述研究结果表明,苏抗103与GK22影响棉田玉米螟幼虫与蛹的存活与生命力,进而影响其成虫繁殖力而表现出良好抗性。     

Parental behaviour in Copris lunaris (Coleoptera, Scarabaeidae): care and defence of brood balls and nest

Abstract. 1. Nesting female beetles righted brood balls (so as to replace the egg or larva in the uppermost position) and repaired damaged balls. This behaviour required the presence of an egg or larva in the ball, or of a short-lasting material found just after oviposition. The shape of the ball was also a righting stimulus since artificial ellipsoids were stood on end.
2. Balls containing dichloromethane extracts of C.lunaris brood were righted and repaired. Eggs and larvae of several other Scarabaeidae did not release these responses but were destroyed.
3. Righting behaviour was released when brood was absent from the top of the ball. The beetle then crawled vertically downwards and, if it encountered the displaced apex, a novel rolling action followed which automatically turned the ball towards the correct position.
4. An opening made in the nest was repaired with soil excavated from the chamber floor. Clunaris adults and Aphodius fossor larvae were attacked if they were encountered in the nest.     

Effect of nutritional condition on larval food requisition behavior in a subterranean termite Reticulitermes speratus (Isoptera: Rhinotermitidae)

Although optimal investment theory would be similarly applicable to eusocial insects to maximize colony reproductive outputs, directly distinguishing an amount of investment in each larva should be a difficult task for workers because of the characteristics of group living. Thus, it is expected that workers adjust brood care by using a cue or signal conveying information of larval status. In termites, which are typical group of eusocial insects, there are nevertheless few direct observations on worker brood care and little is known about cues inducing worker feeding. I show here that a Japanese subterranean termite Reticulitermes speratus uses an overt food solicitation by larva, “pecking”, as a cue for worker feeding. Direct observations demonstrated that workers feed larvae in response to larval pecking. Furthermore, nutritional experiments showed that larvae exhibited pecking more frequently when their nutrient status is lower; hence, pecking may be an honest reflection of larval hunger status. These results indicate that workers can feed more starved larvae than less starved ones because pecking honestly reflects larval hunger state. That is, feeding in response to pecking should standardize the total amount of food intake of each larva and help a termite colony make worker investment efficient.     

Who came first--larvae or adults? origins of bilaterian metazoan larvae

There is a classic controversy in zoology over whether the common ancestor of living bilaterian phyla was a benthic animal with a bilaterian body plan, or was a pelagic larva-like animal similar to what we see today in the primary larvae of indirect-developing bilaterians. We examine the current larva-like adult hypothesis, and present an alternate model for the evolution of complex life histories by intercalation of larval features into the ontogeny of an ancestral direct-developing bilaterian. This gradual accumulation of larval features results in a developmental regulatory program that produces a larva distinct in body plan from the adult. The evolution of a rapid and complete metamorphosis is made possible by the convergent evolution of set aside cells in the final stages of the emergence of indirect developing larval forms. Although convergences abound either hypothesis for the evolution of developmental pathways and life histories, the bilaterian first hypothesis is consistent with all stages of evolution of a complex life history being selectively advantageous, with the rapid evolution of larval forms, and with the frequent co-option of genes from the adult phase of the life cycle prevalent in the evolution of embryos and larvae.     

大蜂螨对一些气味物质的趋性

狄斯瓦螨Varroa destructor Anderson & Trueman是意大利蜜蜂Apis mellifera Spinola的主要外寄生螨。雌成螨在幼虫巢房封盖前不久侵入幼虫巢房,并开始繁殖为害。从雌成螨在一个很短的时间内进入蜜蜂幼虫巢房,以及雄蜂幼虫巢房蜂螨的寄生率明显高于工蜂幼虫巢房的现象,表明蜜蜂幼虫体表一些信息素(semiochemicals)可能起着重要的引诱作用。作者对与大蜂螨相关的19种气味物质进行筛选,并对封盖前工蜂幼虫和雄蜂幼虫表皮挥发物进行气谱及气-质联谱测定。结果表明:雄蜂6龄幼虫对大蜂螨的引诱作用显著高于丁香水等10种气味物质。工蜂和雄蜂末龄幼虫体表挥发物的共有组份是9-二十三烯(C23H46),但它在雄蜂幼虫中所占的比例要明显高于工蜂幼虫。工蜂幼虫的特有主要组分是十八烷(C18H38)和9-甲基十九烷(C19H40);而雄蜂幼虫的特有主要组分是二十五烷(C25H52)和二十三烷(C23H48)。     

A synthesis of genetic connectivity in deep‐sea fauna and implications for marine reserve design

With anthropogenic impacts rapidly advancing into deeper waters, there is growing interest in establishing deep‐sea marine protected areas (MPAs) or reserves. Reserve design depends on estimates of connectivity and scales of dispersal for the taxa of interest. Deep‐sea taxa are hypothesized to disperse greater distances than shallow‐water taxa, which implies that reserves would need to be larger in size and networks could be more widely spaced; however, this paradigm has not been tested. We compiled population genetic studies of deep‐sea fauna and estimated dispersal distances for 51 studies using a method based on isolation‐by‐distance slopes. Estimates of dispersal distance ranged from 0.24 km to 2028 km with a geometric mean of 33.2 km and differed in relation to taxonomic and life‐history factors as well as several study parameters. Dispersal distances were generally greater for fishes than invertebrates with the Mollusca being the least dispersive sampled phylum. Species that are pelagic as adults were more dispersive than those with sessile or sedentary lifestyles. Benthic species from soft‐substrate habitats were generally less dispersive than species from hard substrate, demersal or pelagic habitats. As expected, species with pelagic and/or feeding (planktotrophic) larvae were more dispersive than other larval types. Many of these comparisons were confounded by taxonomic or other life‐history differences (e.g. fishes being more dispersive than invertebrates) making any simple interpretation difficult. Our results provide the first rough estimate of the range of dispersal distances in the deep sea and allow comparisons to shallow‐water assemblages. Overall, dispersal distances were greater for deeper taxa, although the differences were not large (0.3–0.6 orders of magnitude between means), and imbalanced sampling of shallow and deep taxa complicates any simple interpretation. Our analyses suggest the scales of dispersal and connectivity for reserve design in the deep sea might be comparable to or slightly larger than those in shallow water. Deep‐sea reserve design will need to consider the enormous variety of taxa, life histories, hydrodynamics, spatial configuration of habitats and patterns of species distributions. The many caveats of our analyses provide a strong impetus for substantial future efforts to assess connectivity of deep‐sea species from a variety of habitats, taxonomic groups and depth zones.     

Transcriptional signature of accessory cells in the lateral line, using the Tnk1bp1:EGFP transgenic zebrafish line

Background

A metamorphic life-history is present in the majority of animal phyla. This developmental mode is particularly prominent among marine invertebrates with a bentho-planktonic life cycle, where a pelagic larval form transforms into a benthic adult. Metamorphic competence (the stage at which a larva is capable to undergo the metamorphic transformation and settlement) is an important adaptation both ecologically and physiologically. The competence period maintains the larval state until suitable settlement sites are encountered, at which point the larvae settle in response to settlement cues. The mechanistic basis for metamorphosis (the morphogenetic transition from a larva to a juvenile including settlement), i.e. the molecular and cellular processes underlying metamorphosis in marine invertebrate species, is poorly understood. Histamine (HA), a neurotransmitter used for various physiological and developmental functions among animals, has a critical role in sea urchin fertilization and in the induction of metamorphosis. Here we test the premise that HA functions as a developmental modulator of metamorphic competence in the sea urchin Strongylocentrotus purpuratus.

Results

Our results provide strong evidence that HA leads to the acquisition of metamorphic competence in S. purpuratus larvae. Pharmacological analysis of several HA receptor antagonists and an inhibitor of HA synthesis indicates a function of HA in metamorphic competence as well as programmed cell death (PCD) during arm retraction. Furthermore we identified an extensive network of histaminergic neurons in pre-metamorphic and metamorphically competent larvae. Analysis of this network throughout larval development indicates that the maturation of specific neuronal clusters correlates with the acquisition of metamorphic competence. Moreover, histamine receptor antagonist treatment leads to the induction of caspase mediated apoptosis in competent larvae.

Conclusions

We conclude that HA is a modulator of metamorphic competence in S. purpuratus development and hypothesize that HA may have played an important role in the evolution of settlement strategies in echinoids. Our findings provide novel insights into the evolution of HA signalling and its function in one of the most important and widespread life history transitions in the animal kingdom - metamorphosis.