HETEROCHRONIC DEVELOPMENTAL PLASTICITY IN LARVAL SEA URCHINS AND ITS IMPLICATIONS FOR EVOLUTION OF NONFEEDING LARVAE

Preexisting developmental plasticity in feeding larvae may contribute to the evolutionary transition from development with a feeding larva to nonfeeding larval development. Differences in timing of development of larval and juvenile structures (heterochronic shifts) and differences in the size of the larval body (shifts in allocation) were produced in sea urchin larvae exposed to different amounts of food in the laboratory and in the field. The changes in larval form in response to food appear to be adaptive, with increased allocation of growth to the larval apparatus for catching food when food is scarce and earlier allocation to juvenile structures when food is abundant. This phenotypic plasticity among full siblings is similar in direction to the heterochronic evolutionary changes in species that have greater nutrient reserves within the ova and do not depend on particulate planktonic food. This similarity suggests that developmental plasticity that is adaptive for feeding larvae also contributes to correlated and adaptive evolutionary changes in the transition to nonfeeding larval development. If endogenous food supplies have the same effect on morphogenesis as exogenous food supplies, then changes in genes that act during oogenesis to affect nutrient stores may be sufficient to produce correlated adaptive changes in larval development.     

The larvae of Diadema setosum (Leske, 1778) (Camarodonta: Diadematidae) from South China Sea

Abstract

Diadema setosum (Leske, 1778) develops from small isolecithal eggs with a diameter of 84 ± 3 μm. Embryonic development took about 6.5–7 h and finished when a blastula left the fertilization envelope and became a larva. At this stage, the first pigment cells had appeared. At 23 h a prism developed; at 44 h a pluteus with one pair of arms had appeared; at 45 h of development plutei had two pairs of arms. The pigment cells colour the pluteus of D. setosum dark red. When 20-day-old larvae were mechanically stimulated, they flared their arms which may be defensive behaviour. During further development, the post oral arms of plutei grew to 1900 μm or more. Metamorphosis took place at about 40–45 days. At this time, five primary ambulacral podia were visible within the larval body. The duration of metamorphosis from the moment of larval settlement until the juvenile sea urchins began to move along the bottom was 40–60 min. The diameter of the test of the newly metamorphosed juvenile sea urchins was about 500 μm.     

Asymmetric formation and possible function of the primary pore canal in plutei of Temnopleurus hardwicki

The development and possible function of the primary pore canal (PPC) in plutei of the sea urchin Temnopleurus hardwicki was examined by immunochemistry, electron microscopy and microsurgery. Left and right PPC that extended from coelomic sacs in plutei contained a bundle of cilia with a 9 + 2 structure that was initially detected as a group of anti-acetylated tubulin antibody-binding granules in the epithelium of coelomic sacs in 28 h postfertilization (PF) prism larvae. The granules extended to be a bundle of fibers toward the larval dorsal surface, concurrent with formation of the PPC on both sides, over the next 4 h. The cilia in both PPC beat actively. However, the PPC on the right side disappeared by approximately 55 h PF, establishing left-right asymmetry by 60 h PF (the four-arm pluteus stage). The numbers of cilia in the left and right PPC in 56 h PF plutei were five and eight, respectively. Microsurgical removal of the coelomic sac from both sides or the left side only from 26 h PF prism larvae decreased body width to 64 and 91% of normal width by 50 h PF pluteus stage, respectively, whereas that of the right PPC did not. These observations suggest that PPC contribute to the maintenance of normal body width, and that there is asymmetrical activity between the left and right PPC.     

Differential susceptibility of marine invertebrate larvae: Laboratory predation of sand dollar, Dendrasterexcentricus (Eschscholtz), embryos and larvae by zoeae of the red crab, Cancer productus Randall

Predation on eggs, embryos, and larvae of the sand dollar, Dendraster excentricus (Eschscholtz) was investigated in a series of laboratory feeding experiments. Dendraster susceptibility to predation by zoea larvae of the red crab, Cancer productus Randall was strongly dependent on developmental stage and ontogenetic differences in motility. Clearance rates by C. productus were highest for eggs and averaged 0.551·zoea⁻¹·day⁻¹. Embryos and prism larvae of Dendraster were consumed at an intermediate rate, while pluteus larvae were captured at a relatively low rate. Clearance rates decreased from 0.18 to 0.031·zoea⁻¹·day⁻¹ during the transition of prism larvae into echinoplutei. Differences in Dendraster susceptibility to predation cannot be attributed to increasing prey body size because dwarf plutei were captured at the same rate as normal plutei. Reduced capture rates by Cancer productus zoeae are dependent on the development of Dendraster swimming behavior. Periodic reversals in the direction of ciliary beating and backwards swimming effectively remove Dendraster plutei from the immediate capture sphere of Cancer productus. Reversed swimming appears to function as a post-contact encounter response that reduces the mortality rates of Dendraster plutei.     

The sea urchin Lytechinus variegatus lives close to the upper thermal limit for early development in a tropical lagoon

Thermal tolerance shapes organisms' physiological performance and limits their biogeographic ranges. Tropical terrestrial organisms are thought to live very near their upper thermal tolerance limits, and such small thermal safety factors put them at risk from global warming. However, little is known about the thermal tolerances of tropical marine invertebrates, how they vary across different life stages, and how these limits relate to environmental conditions. We tested the tolerance to acute heat stress of five life stages of the tropical sea urchin Lytechinus variegatus collected in the Bahía Almirante, Bocas del Toro, Panama. We also investigated the impact of chronic heat stress on larval development. Fertilization, cleavage, morula development, and 4‐armed larvae tolerated 2‐h exposures to elevated temperatures between 28–32°C. Average critical temperatures (LT₅₀) were lower for initiation of cleavage (33.5°C) and development to morula (32.5°C) than they were for fertilization (34.4°C) or for 4‐armed larvae (34.1°C). LT₅₀ was even higher (34.8°C) for adults exposed to similar acute thermal stress, suggesting that thermal limits measured for adults may not be directly applied to the whole life history. During chronic exposure, larvae had significantly lower survival and reduced growth when reared at temperatures above 30.5°C and did not survive chronic exposures at or above 32.3°C. Environmental monitoring at and near our collection site shows that L. variegatus may already experience temperatures at which larval growth and survival are reduced during the warmest months of the year. A published local climate model further suggests that such damaging warm temperatures will be reached throughout the Bahía Almirante by 2084. Our results highlight that tropical marine invertebrates likely have small thermal safety factors during some stages in their life cycles, and that shallow‐water populations are at particular risk of near future warming.     

Morphogenesis of the digestive tract of the pluteus larva of Strongylocentrotus purpuratus: sphincter formation

In the developing pluteus larva of S. purpuratus, the initial morphogenetic event in the formation of a functional gut is the appearance of two constrictions in the archenteron. These two constrictions become the cardiac and pyloric sphincters. During the 2 h in which the constrictions form, the sphincter cells change from cuboidal to wedge-shaped, and the apical ends of the sphincter cells develop an electron-dense region in which microfilaments can be resolved. Constriction of the archenteron was reversibly inhibited by cytochalasin B, although cytochalasin B had no effect once the constrictions had fully formed. Neither the electron-dense region nor the microfilaments were observed after cytochalasin B treatment. It is suggested that sphincter formation is initially accomplished by a microfilament-mediated contraction of the apical ends of the sphincter cells, which changes their shape and constricts the archenteron.