Mechanistic Basis of Life-History Evolution in Anuran Amphibians: Direct Development

SYNOPSIS. The primitive, or ancestral reproductive mode forRecent amphibians involves a complex, biphasic life history.Yet evolutionarily derived, alternate modes are seen in allthree living orders and predominate in some clades. Analysisof the consequences and mechanistic bases of one such mode—directdevelopment—can provide insights into the evolutionaryopportunities and constraints conferred by the ancestral metamorphicontogeny. Direct development in the anuran genus Eleutherodactylusinvolves fundamental alterations to many features of embryonicand posthatching development. At hatching, young emerge as fullyformed, albeit tiny versions of the adult; most larval featuresare absent. Pervasive changes in ontogenetic timing, in particularthe precocious (embryonic) formation of many adult structures,appear to be correlated with early development of the thyroidaxis, although responsiveness to exogenous thyroid hormone isdiminished or even lacking in at least some peripheral tissues.Changes in cranial patterning are likely mediated by the embryonicneural crest, although many gross features of crest biologyare highly conserved. Laboratory-based analyses of direct developmentand other derived reproductive modes in amphibians, using contemporarymethods developed for more standard, "model" organisms, maycontribute important insights into life-history evolution thatcomplement those derived from analyses of morphology, ecologyand phylogeny.     

Adaptive Strategy and the Origin of Grades and Ground-Plans

The ground-plans of higher metazoans seem to have originatedchiefly in two waves, one near 700 million, the other near 580million years ago. The first wave, involving the origin of thecoelom, was probably associated with invasion of the substrateand the evolution of an infaunal community, while the secondinvolved a reinvasion of the sea-floor surface and the developmentof an epibenthic fauna, for which skeletonization was a commonadaptation. Each of these waves seems to represent adaptationsto patterns of environmental variability—that is, theyoriginate as adaptive strategies. Later waves of diversificationtend to involve lower taxonomic categories but neverthelessappear to have been associated with changes in adaptive strategies.     

The Messenger RNA Population in the Embryonic Axes of Phaseolus vulgaris During Development and Following Germination

Misra, S. and Bewley, J. D. 1985. The messenger RNA populationin the embryonic axes of Phaseolus vulgaris during developmentand following germination.—J. exp. Bot. 36: 1644–1652. Messenger RNAs were extracted from young, mid-maturation, late-maturation,mature-dry, and 20-h-germinated embryonic axes of Phaseolusvulgaris cv. Taylor's Horticultural. They were translated invitro in a rabbit reticulocyte lysate protein synthesizing system.Analysis of the translation products using two-dimensional polyacrylamidegel electrophoresis indicated that there were substantial changesin the messenger RNA populations of developing and germinatingaxes. The number of polypeptides synthesized increased sharplyat 20–22 d after pollination and then declined. Therewas a parallel increase and decrease in the Poly(A)⁺ contentof the seed axis. The analysis showed that certain messageswere present throughout development and were stored in maturedry seed. These messages were degraded upon subsequent rehydration.Some messages appeared during mid-maturation but declined duringlater stages of development and were absent from the matureseed. In the germinating seed a set of messages unique to germinationappeared. Key words: —Seed development, germination, mRNA, in vitro translation, Phaseolus vulgaris     

An Overview of the Organization of the Brain of Actinopterygian Fishes

SYNOPSIS. The brain of actinopterygian fishes can be subdividedinto five principal parts, rhombencephalon, cerebellum, mesencephalon,diencephalon and telencephalon, each of which contains a numberof separate morphological entities: nuclei, areas or zones.Analysis of the origin and termination of the cranial nervesand their components reveals that many of the morphologicalentities distinguished in the actinopterygian brain can be interpretedin terms of elementary sensory and motor functions. Experimentalanatomical and physiological studies on the fiber connectionsof the entities thus defined have led to a functional interpretationof many other parts of the brain. Thus, the central circuitryrelated to such sensory functions as hydrodynamic pressoreception,electroreception, vision, gustation and olfaction are well-known,and the same holds true for the motor systems related to feedingand locomotion. However, there are still many regions in theactinopterygian brain the functional significance of which ispoorly understood, and it should be emphasized that most ofour knowledge on the organization of the brain of this grouprests merely on observations in teleosts. One of the most interestingaspects of actinopterygian neurobiology is that the telencephalonin this group develops in a way which differs fundamentallyfrom that found in all other craniates, and that the telencephalonshows a marked progressive differentiation in the series: polypterids—chondrosteans—halecomorphs—teleosts.     

The evolution of hominoid cranial diversity: A quantitative genetic approach

Hominoid cranial evolution is characterized by substantial phenotypic diversity, yet the cause of this variability has rarely been explored. Quantitative genetic techniques for investigating evolutionary processes underlying morphological divergence are dependent on the availability of good ancestral models, a problem in hominoids where the fossil record is fragmentary and poorly understood. Here, we use a maximum likelihood approach based on a Brownian motion model of evolutionary change to estimate nested hypothetical ancestral forms from 15 extant hominoid taxa. These ancestors were then used to calculate rates of evolution along each branch of a phylogenetic tree using Lande's generalized genetic distance. Our results show that hominoid cranial evolution is characterized by strong stabilizing selection. Only two instances of directional selection were detected; the divergence of Homo from its last common ancestor with Pan, and the divergence of the lesser apes from their last common ancestor with the great apes. In these two cases, selection gradients reconstructed to identify the specific traits undergoing selection indicated that selection on basicranial flexion, cranial vault expansion, and facial retraction characterizes the divergence of Homo, whereas the divergence of the lesser apes was defined by selection on neurocranial size reduction.     

Alternative Concepts of Reproductive Effort, Costs of Reproduction, and Selection in Life-History Evolution

An outline for an organismic theory of reproductive tacticsis presented to develop the demographic theory of optimal reproductivetactics into a more realistic theory of life-history evolution.Reproductive effort—denned as the proportion of resourcesinvested in reproduction—and the costs in somatic investmentdo not automatically result in survival costs. Both the conditionswhere survival costs are produced and the conditions where reproductioncan take place without survival costs are specified. Compensationand threshold hypotheses are put forward to allow weaker correlationsbetween reproduction and survival than the trade-off hypothesis,which assumes direct impacts by reproductive effort on survival.Furthermore, reproductive tactics are unlikely to be mouldedby the demographic forces of selection only. An empirical exampleis shown where residual reproductive value played no significantrole in the evolution of reproductive tactics. Selection probablyoperates not on separate life-history traits but on whole organismsthrough their entire life-history. The structural and physiologicalintercouplings between separate traits can result in phenotypicopportunity sets where selection can mould life-history traitsonly within the constraints of the opportunity sets. Optimizationtheory has provided an efficient technique for modelling andmaking predictions. However, organismic selection does not necessarilyoptimize adaptive strategies but eliminates unfit strategies.Life-history theory, and evolutionary theory in general, canbe developed along alternative logical lines when differenthypotheses are generated on how selection operates.     

How and When Selection Experiments Might Actually be Useful

Laboratory natural selection and artificial selection are vitaltools for addressing specific questions about evolutionary patternsof variation. Laboratory natural selection can illuminate whethera putative selective agent is capable of generating long-term,sustained changes in individual traits and suites of traits.Artificial selection is the essential tool for understandingthe general evolvability of traits and the extent to which geneticcorrelations constrain evolution. We review the contexts inwhich each type of experiment seems capable of offering keyinsights into important evolutionary issues. We also discusstheoretical and methodological considerations that play criticalroles in designing selection experiments that are relevant toevolutionary patterns of trait variation. In particular, wefocus on the critical role of selection intensity and the consequencesof experiments with different intensities. While selection experimentsare not practical in many cases, sophisticated selection experiments—designedwith careful consideration of the theory of selection—shouldbe taken beyond model organisms and used in well-chosen naturalsystems to understand natural patterns of variation.     

Two strategies for investigating the evolution of behavior

In this paper I distinguish and characterize two strategies, both prominent in contemporary biology, for investigating the evolution of behavior. The ‘Lorenzian Strategy’ is taxon-focused, holistic, and particularistic, and relies heavily on naturalistic observation as well as careful experimental manipulation of target systems; it tends to produce detailed knowledge of concrete historical instances of the evolution of behavior in particular lineages. The ‘Analytic Strategy’ is principle-focused, generative, and taxonomically universal; it relies on the development of mathematical principles (simple analytic models) of the evolution of behavior at an abstract level, and uses experimentation to garner support for the empirical relevance for these. The strategies hence employ different methods and produce different sorts of knowledge, hence they are neither inconsistent nor redundant, but complementary, and indeed they both play important roles in the contemporary biology of animal behavior.     

Evolution of Endocrine Regulation of Gastrointestinal Function in Lower Vertebrates

The information available concerning the evolution of endocrineregulation of three gastrointestinal functions in lower vertebrates—gastricacid secretion, gallbladder contraction, and pancreatic enzymesecretion—is reviewed. The actions of hormones of thecholecystokinin/gastrin family of peptides have been the mostwidely studied and are emphasized. It is concluded that regulationof pancreatic enzyme secretion is a primitive action of cholecystokininand that the sensitivity of gallbladder muscle and gastric acid-secretingcells to these peptides evolved later, possibly in the ancestrallineage that led to the gnathostomes. The need for increaseddigestive efficiency to support the higher levels of activitymade possible by the evolution of jaws is suggested as a strongselection pressure leading to this pattern of evolution of endocrineregulation of gastrointestinal function.     

Experimental Evolution and Its Role in Evolutionary Physiology

Four general approaches to the study of evolutionary physiology—phylogenetically-basedcomparisons, genetic analyses and manipulations, phenotypicplasticity and manipulation, and selection studies—areoutlined and discussed. We provide an example of the latter,the application of laboratory selection experiments to the studyof a general issue in environmental adaptation, differencesin adaptive patterns of generalists and specialists. A cloneof the bacterium Escherichia coli that had evolved in a constantenvironment of 37°C was replicated into 6 populations andallowed to reproduce for 2,000 generations in a variable thermalenvironment alternating between 32 and 42°C. As predictedby theory, fitness and efficiency of resource use increasedin this new environment, as did stress resistance. Contraryto predictions, however, fitness and efficiency in the constantancestral environment of 37°C did not decrease, nor didthermal niche breadth or phenotypic plasticity increase. Selectionexperiments can thus provide a valuable approach to testinghypotheses and assumptions about the evolution of functionalcharacters.     

Mechanistic Approaches to Community Ecology: A New Reductionism

Mechanistic approaches to community ecology are those whichemploy individual— ecological concepts—those ofbehavioral ecology, physiological ecology, and ecomorphology—as theoretical bases for understanding community patterns. Suchapproaches, which began explicitly about a decade ago, are justnow coming into prominence. They stand in contrast to more traditionalapproaches, such as MacArthur and Levins (1967),which interpretcommunity ecology almost strictly in terms of "megaparameters.". Mechanistic approaches can be divided into those which use populationdynamics as a major component of the theory and those whichdo not; examples of the two are about equally common. The firstapproach sacrifices a highly detailed representation of individual—ecological processes; the second sacrifices an explicit representationof the abundance and persistence of populations. Three subdisciplines of ecology—individual, populationand community ecology—form a "perfect" hierarchy in Beckner's(1974) sense. Two other subdisciplines—ecosystem ecologyand evolutionary ecology—lie somewhat laterally to thishierarchy. The modelling of community phenomena using sets ofpopulation-dynamical equations is argued as an attempt at explanationvia the reduction of community to population ecology. Much ofthe debate involving Florida State ecologists is over whetheror not such a relationship is additive (or conjunctive), a verystrong form of reduction. I argue that reduction of communityto individual ecology is plausible via a reduction of populationecology to individual ecology. Approaches that derive the population-dynamicalequations used in population and community ecology from individual-ecologicalconsiderations, and which provide a decomposition of megaparametersinto behavioral and physiological parameters, are cited as illustratinghow the reduction might be done. I argue that "sufficient parameters"generally will not enhance theoretical understanding in communityecology. A major advantage of the mechanistic approach is that variationin population and community patterns can be understood as variationin individual-ecological conditions. In addition to enrichingthe theory, this allows the best functional form to be chosenfor modeling higher-level phenomena, where "best" is definedas biologically most appropriate rather than mathematicallymost convenient. Disadvantages of the mechanistic approach arethat it may portend an overly complex, massive and special theory,and that it naturally tends to avoid many-species phenomenasuch as indirect effects. The paper ends with a scenario fora mechanistic-ecological utopia.     

Molecular Variation and Possible Lines Of Evolution of Peptide and Protein Hormones

The widespread distribution of certain steroids and amino acidderivatives with hormonal properties is considered evidencein support of the dictum that "it is not the hormones that change,but rather the uses to which they are put." However, analysesof the distributions, biological activities, immunological cross-reactivities,and sequences of amino acids of five representative peptideand protein hormones or groups of hormones—lactogenichormone, growth hormone, the corticotropin-MSH-ß lipotropinfamily, insulin, and the neurohypophysial hormones—supporta concept of change and of molecular evolution of these polypeptidicmolecules. When analyzed in terms of the genetic code, the aminoacid interchanges which have been revealed by determinationof sequences of amino acids can, most often, be explained bysingle base mutations in the appropriate codons. In two instanceswhere two base mutations within a single codon are required,intermediate replacements of amino acid have been suggested;one of these would lead to a 2-ALA-ß MSH, and theother to a 4-PRO, 8-ILE oxytocin.     

Histology of Callogenesis and Somatic Embryogenesis Induced in Stem Fragments of Cork Oak (Quercus suber) Cultured In Vitro

Calluses able to produce somatic embryos were formed duringin vitro culture of shoot fragments of cork oak (Quercus suberL.).Histological monitoring of these fragments during cultureshowed that it was the cortical parenchyma cells which underwentdedifferentiation before calluses were formed by repeated divisions.The calluses consisted of parenchyma cells surrounded by a fewlayers of meristematic cells. Proembryos formed in groups aroundthe edge of some calluses. Histological examination showed thatthey were produced by the evolution of two different categoriesof cell: one category had the appearance of ‘embryogenic’cells with very thick walls, a small vacuole rich in starchand a well-developed nucleus with a prominent nucleolus. Theother cells were very bulky with large vacuoles; their morphologywas similar to that of suspensor cells encountered in embryogenesisin gymnosperms. The ontogenic stages were similar to those describedin zygotic embryos of the genus Quercus. Nevertheless, mostof the embryonic structures deviated from normal developmentand at all stages produced secondary proembryos. Cork-oak, Quercus suber L, histology, callogenesis, somatic embryogenesis, embryogenic cells, starch, secondary embryogenesis     

Monte Carlo simulation program for ecosystems

A Monte Carlo computer simulation program is designed in orderto describe the spatial and time evolution of a population ofliving individuals under preassigned environmental conditionsof energy. The simulation is inspired by previous techniquesdeveloped in physics — in particular, in molecular dynamicsand simulations of liquids — and it already provides somenew insights regarding macroscopic deterministic models in ecologyand concerning eventual control of artificial biomass productionplants. Received on July 15, 1986; accepted on October 9, 1986     

Ontogenesis and Morphological Diversification

SYNOPSIS. The role of development in constraining the directionalityand patterns of morphological evolution is examined. The natureof morphological variation and appearance of morphological noveltiesis determined by the epigenetic properties of the organism.Consideration of these properties has profound implicationsfor current theories of morphological evolution. Developmentalconstraints impose severe limitations on the gradualistic actionof directional selection. Evolutionis viewed as the result ofdifferential survival of morphological novelties. However, theproduction of morphological novelties by developmental programsis not random. This non-randomness in morphologically expressedgenetic mutations—an epigenetic property—can resultin phyletic trends, parallelisms and convergences.     

Chnage, regularity, and value in the evolution of animal learning

I present a simple model that considers how three factors—change,regularity, and value— influence the evolution of animallearning. Change and regularity are considered by introducingtwo terms that measure environmental persistence. One term,"between-generation persistence, " defines the extent to whichstates in the parental generation predict states in the offspringgeneration; the other term, "within-generation persistence,"defines the extent to which today predicts tomorrow within anindividual's lifetime. Within-generation persistence is shownto be the most important of these two terms. When there is somechange, increasing the within-generation persistence promotesthe evolution of learning, and the between-generation persistenceterm has no effect. However, when the environment is almostcompletely fixed, then increasing change, either within or betweengenerations, promotes the evolution of learning. This occursbecause (1) the change required to promote the evolution oflearning can occur either within or between generations eventhough (2) the regularity required to promote the evolutionof learning must come within an animal's lifetime. The regionof absolute fixity, in which learning does not generally evolve,is relatively small. The results for value, or payoffs, suggestthat learning is most useful when all the alternatives to learningyield about the same payoff. [Behav Ecol 1991; 2: 77–89]     

Reproductive Development and Nitrogen Fixation in Soybean (Glycine max [L.] Merril)

Gomes, M. A. F. and Sodek, L. 1987. Reproductive developmentand nitrogen fixation in soybean (Glycine max (L.) Merril).—J.exp. Bot. 38: 1982–1987. Nitrogenase activity (acetylene reduction) was measured duringthe growth cycle of soybean plants induced to flower at twodifferent ages. The decline in nitrogenase activity towardsthe end of the cycle was clearly associated with pod-fillingfor both flowering dates when plants were cultivated under lowerlight and temperature conditions (out of season). Under higherlight and temperature conditions (normal growing season) thedecline was independent of the flowering date. Furthermore,the timing of the decline was not altered when plants were maintainedunder long-day (vegetative) conditions nor when flowers wereremoved. It is suggested that under more favourable growth conditionsthe diversion of assimilates by the fruits is not the primarycause of the decline in nodule activity, but competition bythe fruits may be important when the production of photo-assimilatesis more limited. Key words: Glycine max, nitrogenase, source-sink     

Cranial integration in Homo: singular warps analysis of the midsagittal plane in ontogeny and evolution

This study addresses some enduring issues of ontogenetic and evolutionary integration in the form of the hominid cranium. Our sample consists of 38 crania: 20 modern adult Homo sapiens, 14 sub-adult H. sapiens, and four archaic Homo. All specimens were CT-scanned except for two infant H. sapiens, who were imaged by MR instead. For each specimen 84 landmarks and semi-landmarks were located on the midsagittal plane and converted to Procrustes shape coordinates. Integration was quantified by the method of singular warps, a new geometric-statistical approach to visualizing correlations among regions. The two classic patterns of integration, evolutionary and ontogenetic, were jointly explored by comparing analyses of overlapping subsamples that span ranges of different hypothetical factors. Evolutionary integration is expressed in the subsample of 24 adult Homo, and ontogenetic integration in the subsample of 34 H. sapiens. In this data set, vault, cranial base, and face show striking and localized patterns of covariation over ontogeny, similar but not identical to the patterns seen over evolution. The principal differences between ontogeny and phylogeny pertain to the cranial base. There is also a component of cranial length to height ratio not reducible to either process. Our methodology allows a separation of these independent processes (and their impact on cranial shape) that conventional methods have not found.     

Olfaction in Mammals

This paper centers on selected—and particularly, littlerecognized— problems in mammalian olfaction: (1) Withcertain exceptions the spacing of the external nares in mostmammals does not favor orientation in an odor gradient by simultaneouscomparisons of odor intensities (tropotaxis). (2) The mammaliannose is rich in both dynamic and static devices for conditioningand controlling the How of inspired air. (3) A well-developedvomeronasal organ is widely distributed but its function isobscure. (4) Nerve impulse traffic telemetered from the olfactorybulb of freely moving rats shows a varying pattern of discretebursts of units with each inspiration and more sustained discharges.(5) Olfaction in species showing adaptations for lite in water,air, and underground is reviewed. (6) Because of its rich olfactory-trigeminalinnervation the snout of pigs, moles, etc., may be consideredas a "chemotactile" organ.