Shrinking to fit: fluid jettison from a haemocoelic hydrostatic skeleton during defensive withdrawals of a gastropod larva

Although most of the basic animal body plans are supported by hydrostatic skeletons consisting of fluid maintained at constant volume, studies on how animals have solved biomechanical scaling dilemmas during evolution of large body size have emphasized cases where skeletons are formed by rigid solids. Larvae of gastropod molluscs swim using ciliated velar lobes supported by a constant volume hydrostatic skeleton. Defensive behaviour involves rapid withdrawal of the velar lobes and foot into a protective biomineralized shell. Some gastropod larvae grow to giant size and the velar lobes enlarge allometrically, but the lobes and foot of many can still withdraw completely into the mineral-stiffened shell. I dyed internal fluid of a large gastropod larva with fluorescein to show that fluid supporting the extended velar lobes is expelled from discrete release sites during defensive withdrawals. Scanning electron microscopy suggested that release sites are distinctive papillae on the upper velar epidermis. Ultrathin sections revealed that branched tracks of microvilli-free membrane on the surface of these papillae were formed by very thin epithelial cells, which may rupture and re-anneal during and after defensive withdrawals. Behaviours facilitated by fluid discharge from a haemocoelic (non-coelomic) body compartment have been rarely reported among aquatic invertebrates, but may be more widespread than currently recognized.     

Testing mechanistic models of growth in insects

Insects are typified by their small size, large numbers, impressive reproductive output and rapid growth. However, insect growth is not simply rapid; rather, insects follow a qualitatively distinct trajectory to many other animals. Here we present a mechanistic growth model for insects and show that increasing specific assimilation during the growth phase can explain the near-exponential growth trajectory of insects. The presented model is tested against growth data on 50 insects, and compared against other mechanistic growth models. Unlike the other mechanistic models, our growth model predicts energy reserves per biomass to increase with age, which implies a higher production efficiency and energy density of biomass in later instars. These predictions are tested against data compiled from the literature whereby it is confirmed that insects increase their production efficiency (by 24 percentage points) and energy density (by 4 J mg⁻¹) between hatching and the attainment of full size. The model suggests that insects achieve greater production efficiencies and enhanced growth rates by increasing specific assimilation and increasing energy reserves per biomass, which are less costly to maintain than structural biomass. Our findings illustrate how the explanatory and predictive power of mechanistic growth models comes from their grounding in underlying biological processes.     

The shape of things to come: linking developmental plasticity to post‐metamorphic morphology in anurans

Development consists of growth and differentiation, which can be partially decoupled and can be affected by environmental factors to different extents. In amphibians, variation in the larval environment influences development and causes changes in post‐metamorphic shape. We examined post‐metamorphic consequences, both morphological and locomotory, of alterations in growth and development. We reared tadpoles of two phylogenetically and ecologically distant frog species (the red‐eyed treefrog Agalychnis callidryas and the African clawed frog Xenopus laevis) under different temperatures with ad libitum food supply and under different food levels at a constant temperature. Low temperature and low food levels both resulted in similarly extended larval periods. However, low temperature yielded relatively long‐legged frogs with a lower degree of ossification than warm temperature, whereas low food yielded relatively short‐legged frogs with a higher degree of ossification than high food levels. Such allometric differences had no effect on locomotor performance of juveniles. Our results provide a basis for understanding the relationship between growth, differentiation and post‐metamorphic shape in anurans and help explain many of the discrepancies reported in previous studies.     

Comparative limb bone scaling in turtles: Phylogenetic transitions with changes in functional demands?

Several terrestrial vertebrate clades include lineages that have evolved nearly exclusive use of aquatic habitats. In many cases, such transitions are associated with the evolution of flattened limbs that are used to swim via dorsoventral flapping. Such changes in shape may have been facilitated by changes in limb bone loading in novel aquatic environments. Studies on limb bone loading in turtles found that torsion is high relative to bending loads on land, but reduced compared to bending during aquatic rowing. Release from torsion among rowers could have facilitated the evolution of hydrodynamically advantageous flattened limbs among aquatic species. Because rowing is regarded as an intermediate locomotor stage between walking and flapping, rowing species might show limb bone flattening intermediate between the tubular shapes of walkers and the flattened shapes of flappers. We collected measurements of humeri and femora from specimens representing four functionally divergent turtle clades: sea turtles (marine flappers), softshells (specialized freshwater rowers), emydids (generalist semiaquatic rowers), and tortoises (terrestrial walkers). Patterns of limb bone scaling with size were compared across lineages using phylogenetic comparative methods. Although rowing taxa did not show the intermediate scaling patterns we predicted, our data provide other functional insights. For example, flattening of sea turtle humeri was associated with positive allometry (relative to body mass) for the limb bone diameter perpendicular to the flexion-extension plane of the elbow. Moreover, softshell limb bones exhibit positive allometry of femoral diameters relative to body mass, potentially helping them maintain their typical benthic position in water by providing additional weight to compensate for shell reduction. Tortoise limb bones showed positive allometry of diameters, as well as long humeri, relative to body mass, potentially reflecting specializations for resisting loads associated with digging. Overall, scaling patterns of many turtle lineages appear to correlate with distinctive behaviors or locomotor habits.     

东灵山不同林型五角枫叶性状异速生长关系随发育阶段的变化

分析植物叶片性状种内水平的异速生长关系有助于加深理解生长发育过程中叶片的资源利用模式.分析了东灵山3个主要林型(白桦林、胡桃楸林、辽东栎林)中五角枫成树、幼树、幼苗的叶面积、叶体积、叶含水量与叶干重之间的异速生长关系.结果表明:成树叶面积增长速度小于叶干重的增长速度,幼树、幼苗叶面积与叶干重保持同速增长;成树、幼树叶干重与叶体积保持同速增长,幼苗叶体积的增长速度超过了干重的增长速度;成树叶含水量的增长速度小于叶干重的增长速度,幼树、幼苗两性状间保持等速增长.叶含水量与叶干重的异速生长指数在不同的林型间有显著差异,白桦林叶含水量的增长速度小于叶干重的增长速度,其余两个林型均为等速增长趋势.这些结果揭示了不同发育阶段五角枫资源利用方式的转变.随叶干重的增加,成树将更多的叶生物量投资于同化和支持结构;幼树则保持对光合面积和光合同化结构的稳步投资;而幼苗主要投资于叶面积的增大.叶含水量与叶干重的异速曲线在不同林型间的差异说明叶片代谢活性相较于其他叶性状可塑性更高.     

Phenotypic plasticity of growth trajectory and ontogenic allometry in response to density for eucalyptus hybrid clones and families

BACKGROUND: and Aims Response to density is a crucial aspect of the ecology of trees in forests and plantations. Few studies have investigated the genetics of plasticity in response to density for growth traits such as height and circumference through development. METHODS: Two experiments were carried out in the field, the first with full-sib families of Eucalyptus urophylla x E. grandis hybrids, and the second with clones of E. tereticornis x E. grandis hybrids planted across a range of densities (625, 1111 and 2500 trees ha-1). Height, circumference and stem taper were measured through development in both experiments. Variance components were estimated and a repeated measure approach for plasticity and three different methods were used to compare the variance-covariance matrix across densities. KEY RESULTS: Genetic variance was significantly different from zero but the density x genotype interaction was significant only for clone experiments at the adult stage. Significant plasticity for three traits in both experiments was found. In the clone experiments, a significant clone x time x density interaction was found, suggesting that plasticity for growth and stem form is under genetic control. In both experiments, density did not affect environmental correlation, which remained high throughout tree development. The impact of density on genetic correlation was marked in the clone experiment, with a reduced value at lower density, but was not observed in the family trial. The differences between clones and family are mainly explained by the distribution of genetic variation within and among genotypes. CONCLUSIONS: The results suggest that plasticity for growth traits and form of tropical Eucalyptus species is under genetic control and that the environment changes genetic co-variation through ontogeny. The findings confirm that a tree population with a narrow genetic basis (represented by clones) is sensitive to a changing environment, whereas a population with a broader genetic basis (full-sib family here) exhibits a more stable reaction.