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.     

Mutual support and selection between branches of damaged plants

Snow (1931) showed that the fate of a branch of a plant that had suffered local herbivory could be determined by correlative effects of other branches. This neglected work was continued in the current study, herbivory being simulated by the removal of leaves of different ages from pea plants with two branches. A damaged branch was suppressed when an undamaged alternative branch was present; otherwise the damage never prevented continued development. The removal of mature leaves had a smaller effect than the removal of immature, expanding leaves. When leaves were removed from both branches it was the branches that suffered less damage to their immature leaves that continued shoot development. Branches from which all photosynthetic leaves were repeatedly removed developed only when they retained their immature leaves and remained dominant, inhibiting the development — but not the photosynthesis — of the other branch on the same plant. Accounting for these results requires mechanisms that compare the different branches of the plant and select for development the ones that have the greatest potential for future, rather than present, photosynthesis. It is concluded that a compartmentalized or sectorial distribution of essential substrates can be modified by correlative relations that are probably mediated by hormones.     

Protein structural plasticity exemplified by insertion and deletion mutants in T4 lysozyme.

To further investigate the ways in which proteins respond to changes in the length of the polypeptide chain, a series of 32 insertions and five deletions were made within nine different alpha-helices of T4 lysozyme. In most cases, the inserted amino acid was a single alanine, although in some instances up to four residues, not necessarily alanine, were used. Different insertions destabilized the protein by different amounts, ranging from approximately 1 to 6 kcal/mol. In one case, no protein could be obtained. An "extension" mutant in which the carboxy terminus of the molecule was extended by four alanines increased stability by 0.3 kcal/mol. For the deletions, the loss in stability ranged from approximately 3 to 5 kcal/mol. The structures of six insertion mutants, as well as one deletion mutant and the extension mutant, were determined, three in crystal forms nonisomorphous with wild type. In all cases, including previously described insertion mutants within a single alpha-helix, there appears to be a strong tendency to preserve the helix by translocating residues so that the effects of the insertion are propagated into a bend or loop at one end or the other of the helix. In three mutants, even the hydrophobic core was disrupted so as to permit the preservation of the alpha-helix containing the insertion. Translocation (or "register shift") was also observed for the deletion mutant, in this case a loop at the end of the helix being shortened. In general, when translocation occurs, the reduction in stability is only moderate, averaging 2.5 kcal/mol. Only in the most extreme cases does "bulging" or "looping-out" occur within the body of an alpha-helix, in which case the destabilization is substantial, averaging 4.9 kcal/mol. Looping-out can occur for insertions close to the end of a helix, in which case the destabilization is less severe, averaging 2.6 kcal/mol. Mutant A73-[AAA] as well as mutants R119-[A] and V131-[A], include shifts in the backbone of 3-6 A, extending over 20 residues or more. As a result, residues 114-142, which form a "cap" on the carboxy-terminal domain, undergo substantial reorganizations such that the interface between this "cap" and the rest of the protein is altered substantially. In the case of mutant A73-[AAA], two nearby alpha-helices, which form a bend of approximately 105 degrees in the wild-type structure, reorganize in the mutant structure to form a single, essentially straight helix. These structural responses to mutation demonstrate the plasticity of protein structures and illustrate ways in which their three-dimensional structures might changes during evolution.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Variation in growth and age at maturity in bluegill sunfish: genetic or environmental effects?

Populations of bluegill sunfish Lepomis macrochirus , experiencing heavy juvenile predation, showed increased growth rates and increased age and size at maturity relative to populations experiencing decreased predation on juveniles but increased predation on adults. This study examined bluegills experimentally from both types of populations and a cross between them in a common environment to determine if variation in growth and age at maturity is genetically or environmentally induced. Two factorial experiments, varying strain of bluegills and resource availability, were used to evaluate differences in growth rate. One experiment, varying strain of bluegills, was used to assess differences in age at maturity. Growth was strongly influenced by resource level, but growth rate did not vary among populations. Nearly all bluegills in each population matured at 1 year of age in a common environment. Thus, variation observed in source populations must be mostly attributable to differences in the environment between populations. At least three factors could potentially cause differences in growth and age at maturity: (1) variation in resource availability; (2) variation in demographic structure; and (3) variation in size-specific mortality rates caused by differences in predator abundance between populations. Observed patterns of variation between populations are best explained by effects of differences in predator populations.     

Phylogeny and adaptive radiation within the Anomopoda: a preliminary exploration

The distinctness of the Anomopoda and the polyphyletic nature of the so-called Cladocera are emphasized.An attempt is made to reconstruct the ancestral anomopod, which probably lived in Palaeozoic times. This task is facilitated by the availability of detailed information on extant forms, which includes functional as well as purely morphological considerations and enables us to understand the means whereby complex mechanisms were transformed during evolution. Comparative studies on the ecology and habits of extant forms also throw light on the probable way of life of the ancestral anomopod.Adaptive radiation within the Anomopoda is briefly surveyed and an outline of the suggested phylogeny of the order is indicated.Institute of Environmental and Biological Sciences, University of Lancaster     

Islands: stability, diversity, conservation

Islands present both a diversity and a stability paradox. They are often highly species-poor but have considerable biological interest in terms of extraordinary endemic genera and taxonomically isolated groups. They appear to be stable, as in some cases these organisms have persisted for many millions of years, and having an oceanic climate, extreme climatic events may be comparatively rare. However, when subject to extrinsic (anthropogenic) disturbance they do not appear to be stable, but often suffer catastrophic ecological change. These apparent paradoxes are resolved when it is realized that all these features are consequences of the same island characteristics: biotic isolation and oceanicity. As a result of these two characteristics, far oceanic islands are quantitatively different from continental systems in the nature of their ecological processes, which appear to give rise to an extreme punctuated equilibrium model of evolutionary change. Endemics may be ancient relict endemics displaying prolonged stasis and persistence, or products of adaptive radiation representing rapid punctuational events. A process-based definition of a relict endemic (palaeoendemic) is one whose founding lineage (i.e. the original continental source taxon) has not left any descendents. A corollary of this definition is that the time of divergence between an endemic and its continental sister-group should predate the colonization of the island by the now endemic lineage. An example is Dicksonia arborescens which has been on St Helena for at least 9 Myrs and no longer occurs in the likely source area of Africa. These relict endemics, frequent on islands, are important as the last remnants of tranches of biodiversity that have vanished elsewhere. Island conservation strategies require an integrated understanding of both sides of the diversity and stability paradox so that both island processes and island organisms can be conserved.     

Regulation of neuropeptide expression in the brain by neurotrophins

Neurotrophins, which are structurally related to nerve growth factor, have been shown to promote survival of various neurons. Recently, we found a novel activity of a neurotrophin in the brain: Brain-derived neurotrophic factor (BDNF) enhances expression of various neuropeptides. The neuropeptide differentiation activity was then compared among neurotrophins both in vivo and in vitro. In cultured neocortical neurons, BDNF and neurotrophin-5 (NT-5) remarkably increased levels of neuropeptide Y and somatostatin, and neurotrophin-3 (NT-3) also increased these peptides but required higher concentrations. At elevating substance P, however, NT-3 was as potent as BDNF. In contrast, NGF had negligible or no effect. Neurotrophins administered into neonatal brain exhibited slightly different potencies for increasing these neuropeptides: The most marked increase in neuropeptide Y levels was obtained in the neocortex by NT-5, whereas in the striatum and hippocampus by BDNF, although all three neurotrophins increased somatostatin similarly in all the brain regions examined. Overall spatial patterns of the neuropeptide induction were similar among the neurotrophins. Neurons in adult rat brain can also react with the neurotrophins and alter neuropeptide expression in a slightly different fashion. Excitatory neuronal activity and hormones are known to change expression of neurotrophins. Therefore, neurotrophins, neuronal activity, and hormones influence each other and all regulate neurotransmitter/peptide expression in developing and mature brain. Physiological implication of the neurotransmitter/peptide differentiation activities is also discussed.     

Neuropeptide messenger plasticity in the CNS neurons following axotomy

Neuronal peptides exert neurohormonal and neurotransmitter (neuromodulator) functions in the central nervous system (CNS). Besides these functions, a group of neuropeptides may have a capacity to create cell proliferation, growth, and survival. Axotomy induces transient (1–21 d) upregulation of synthesis and gene expression of neuropeptides, such as galanin, corticotropin releasing factor, dynorphin, calcitonin gene-related peptide, vasoactive intestinal polypeptide, cholecystokinin, angiotensin II, and neuropeptide Y. These neuropeptides are colocalized with “classic” neurotransmitters (acetylcholine, aspartate, glutamate) or neurohormones (vasopressin, oxytocin) that are downregulated by axotomy in the same neuronal cells. It is more likely that neuronal cells, in response to axotomy, increase expression of neuropeptides that promote their survival and regeneration, and may downregulate substances related to their transmitter or secretory activities.     

Genetic differentiation in Plantago major: Ca2+- and Mg2+-stimulated ATPases from roots and their role in phenotypic adaptation

The Mg²⁺- and Ca²⁺-stimulated ATPase activities of the microsomal fractions of the roots of four inbred lines of Plantago major L. were followed at two levels of mineral nutrition. In addition the response of a transfer of plants from one condition to the other was studied. Kinetic properties of the ATPases (K_m and V_max) were calculated and used to differentiate between genetic differences among the inbred lines and the plasticity within each inbred line. The V_max values of the ATPase activity differed significantly between the lines and were directly related to seed number per capsule (low V_max→ 11 seeds per capsule, high V_max→ 33 seeds per capsule). In addition, the V_max values of the ATPase acitivty may be related to ecological strategy. Plasticity of enzyme activity is expressed in differences in the V_max values of the ATPase activity, as a response to nutritional level or changes of the strength of the nutrient solution. Differences in this plasticity in the four selected lines and in rapidity of response to a change in mineral nutrition were directly related to the ecological strategy. These results are discussed in relation to the strategy of the genotypes for survival in the field. The presence of plasticity in line 4 (ssp. pleiospema ) makes this genotype behave like an annual plant, following a ruderal strategy. The absence of plasticity in line 1 (ssp. major ) fits a more competitive strategy.