BackgroundA 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.ResultsOur 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.ConclusionsWe 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.

Ambient calcium is declining in thousands of soft-water lake habitats in temperate regions as a consequence of unsustainable forestry practices, decreased atmospheric calcium deposition and acidic deposition. As their exoskeleton is heavily reinforced with calcium, freshwater crustaceans have a high specific calcium requirement relative to other aquatic organisms. Daphnia, in particular, is an ideal crustacean for investigating the consequences of calcium decline because it is an abundant and important member of freshwater zooplankton communities. Although it has been established that adult and juvenile Daphnia have different tolerances to low ambient calcium as a result of their different life stage-specific calcium requirements, the consequences of declining calcium on embryonic development have never been investigated. Here, we describe the distribution of calcium in embryonic stages of D. magna and introduce a novel and easy to use staging scheme. We tested whether calcium can be traced from mothers to their offspring. Finally, we assessed the fitness consequences of maternal provisioning in limiting calcium environments. We found that while embryos require calcium for their development and moulting, they do not equilibrate with environmental calcium levels. Instead, we were able to trace calcium from mothers to their offspring. Furthermore, our data strongly suggest that females are faced with an allocation trade-off between providing calcium to their offspring and using it for growth and moulting. Together, these data provide novel insights into the consequences of calcium decline for freshwater zooplankton.

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