Culturing echinoderm larvae through metamorphosis.

Echinoderms are favored study organisms not only in cell and developmental biology, but also physiology, larval biology, benthic ecology, population biology and paleontology, among other fields. However, many echinoderm embryology labs are not well-equipped to continue to rear the post-embryonic stages that result. This is unfortunate, as such labs are thus unable to address many intriguing biological phenomena, related to their own cell and developmental biology studies, that emerge during larval and juvenile stages. To facilitate broader studies of post-embryonic echinoderms, we provide here our collective experience rearing these organisms, with suggestions to try and pitfalls to avoid. Furthermore, we present information on rearing larvae from small laboratory to large aquaculture scales. Finally, we review taxon-specific approaches to larval rearing through metamorphosis in each of the four most commonly-studied echinoderm classes-asteroids, echinoids, holothuroids and ophiuroids.

Relationships among egg size, composition, and energy: a comparative study of geminate sea urchins.

Egg size is one of the fundamental parameters in the life histories of marine organisms. However, few studies have examined the relationships among egg size, composition, and energetic content in a phylogenetically controlled context. We investigated the associations among egg size, composition, and energy using a comparative system, geminate species formed by the closure of the Central American Seaway. We examined western Atlantic (WA) and eastern Pacific (EP) species in three echinoid genera, Echinometra, Eucidaris, and Diadema. In the genus with the largest difference in egg size between geminates (Echinometra), the eggs of WA species were larger, lipid rich and protein poor compared to the smaller eggs of their EP geminate. In addition, the larger WA eggs had significantly greater total egg energy and summed biochemical constituents yet significantly lower egg energy density (energy-per-unit-volume). However, the genera with smaller (Eucidaris) or no (Diadema) differences in egg size were not significantly different in summed biochemical constituents, total egg energy, or energy density. Theoretical models generally assume a strong tradeoff between egg size and fecundity that limits energetic investment and constrains life history evolution. We show that even among closely-related taxa, large eggs cannot be assumed to be scaled-up small eggs either in terms of energy or composition. Although our data comes exclusively from echinoid echinoderms, this pattern may be generalizable to other marine invertebrate taxa. Because egg composition and egg size do not necessarily evolve in lockstep, selective factors such as sperm limitation could act on egg volume without necessarily affecting maternal or larval energetics.

Egg size as a life history character of marine invertebrates: Is it all it's cracked up to be?

Egg size is one of the most important aspects of the life history of free-spawning marine organisms, and it is correlated with larval developmental mode and many other life-history characters. Egg size is simple to measure and data are available for a wide range of taxa, but we have a limited understanding of how large and small eggs differ in composition; size is not always the best measure of the characters under selection. Large eggs are generally considered to reflect increased maternal investment, but egg size alone can be a poor predictor of energetic content within and among taxa. We review techniques that have been used to measure the energetic content and biochemical makeup of invertebrate eggs and point out the strengths and difficulties associated with each. We also suggest a number of comparative and descriptive approaches to biochemical constituent analysis that would strengthen our understanding of how natural selection shapes oogenic strategies. Finally, we highlight recent empirical research on the intrinsic factors that drive intraspecific variation in egg size. We also highlight the relative paucity of these data in the literature and provide some suggestions for future research directions.

Evolutionary responses to environmental heterogeneity in central american echinoid larvae: plastic versus constant phenotypes.

Do changes in food resources lead to evolutionary changes in phenotypic plasticity or in different constant phenotypes? I addressed this question by studying plasticity of larval feeding arms for "geminate species pairs" in three echinoid genera. These closely related species were geographically isolated when the Panamanian Isthmus raised 2.8-3.1 million years ago, creating two different food level environments: high but variable food levels in the eastern Pacific versus chronically low food levels in the western Caribbean. I reared larvae of geminate species in different replicated food environments for 10 days postfertilization, collected morphological measurements of individual arm and body lengths, and calculated degrees of plasticity of relative arm length for each species. In contrast to previous studies with temperate echinoids, there was no significant plasticity of arm length in either the Pacific or Caribbean species considered here. Caribbean species, however, had significantly longer relative arm lengths than Pacific species, regardless of food levels. These results suggest that historical changes in food levels have led to the evolution of constant rather than plastic differences between Pacific and Caribbean echinoids. The evolution of plasticity may be limited by the timing of reproduction or by egg size in this system.

Developmental plasticity in Macrophiothrix brittlestars: are morphologically convergent larvae also convergently plastic?

The pluteus larval forms of sea urchins (echinoids) and brittlestars (ophiuroids) use an internal skeleton to project arms that bear a long ciliated band used in swimming and feeding. The length of this ciliated band influences rates of maximum food clearance for larvae of both echinoderm classes and affects rates of growth and development in the plankton. Phylogenetic and morphological evidence, however, tend to support the view that the pluteus morphologies of the two classes are independently derived. Studies with echinoplutei have shown that investment in skeletal growth and ciliated band length changes in response to food conditions, with poorly fed larvae investing more in growth of the larval skeleton and arms either absolutely or in relation to other larval or developing postlarval structures. We present evidence for similar plasticity of skeletal growth in ophioplutei. We examined four species in the brittlestar genus Macrophiothrix that spanned a 3.8-fold range in egg size. Sibling larvae in 14 male-female crosses were reared with high (H) or low (L) food rations, and measurements were recorded for five skeletal arm rods and three non-arm body dimensions. The expression of adaptive plasticity (significantly longer arms in L versus H cultures on a given day) was apparent for most crosses in M. koehleri, the species with the smallest egg size. In the single cross for M. longipeda, larvae from L cultures had longer arms for their body length or stomach width than did larvae from H cultures. In these cases, plasticity was similar in timing, persistence, and magnitude to previously published results from echinoplutei. If internal skeletons are independently derived in the two classes, then plasticity in the expression of this homoplastic trait may itself be homoplastic.

Fisheries: mislabelling of a depleted reef fish.

Any fish species that appears to be readily available in the marketplace will create an impression among the public that there is a plentiful supply of that fish in the sea, but this may belie the true state of the fisheries' stock. Here we use molecular genetic analysis to show that some three-quarters of the fish sold in the United States as 'red snapper'--the US Food and Drug Administration's legally designated common name for Lutjanus campechanus--belong to another species. Mislabelling to this extent not only defrauds consumers but could also adversely affect estimates of stock size if it influences the reporting of catch data that are used in fisheries management.