The role of heterochrony in the establishment of body plan in higher echinoderm taxa

The analysis based on paleontological data shows that the body plans of higher echinoderm taxa were established through the combination of previously developed characters. These combinations appeared due to various heterochronies and resulted in more or less complete filling of the morphological space of logical capabilities. The maximum rank of new taxa decreased with time. New body plans of higher taxa did not replace the old plans but rather overlay them, extending the hierarchy of body plans and the respective hierarchy of taxa. The macroevolution of echinoderms and other metazoans progressed from the formation of an archetype (a general body plan) to individual details, the development of structural plans of lower levels. Heterochrony resulted in mosaic evolution and obscurity of intermediate forms.     

Process heterochronies in endochondral ossification

Heterochrony, evolutionary changes in developmental rates and timing, is a key concept in the construction of a synthesis of development and evolution. Heterochronic changes in vertebrate evolution have traditionally been identified through plesiomorphic-apomorphic comparisons of bone growth. This methodological framework assumes that observed heterochronies are the outcome of dissociations of developmental processes in time. Recent findings of non-heterochronic developmental changes underlying morphological heterochrony invalidate this assumption. In this paper, a function for bone growth (at the organ level) has been mathematically deduced from the underlying developmental mechanisms. The temporal domain of the model spans from the time at maximum growth rate, after the formation of growth plates, to the time at atrophy of the proliferating stratum of cells. Three organizational levels were considered: (a) cell kinetics of endochondral ossification, (b) variation of bone growth rates and (c) variation of accumulated bone growth with increasing age. This quantitative model provides an excellent tool to deal with the problem of the developmental basis of morphological change. I have modelled potential evolutionary changes on the system at different levels of biological organization. This new framework involves an epistemological shift in heterochronic analysis from a pattern-oriented inductive way to a process-oriented deductive way. The analysis of the relationships between the evolutionary alterations of endochondral ossification and the morphological expression of these changes reveals that observed pattern heterochronies can be the outcome of different process heterochronies. Moreover, I discuss at length the heteroposic hypothesis, that evolutionary changes in the tight regulation of the amount of protein synthesized by a cell population during development would underlie acceleration or deceleration in cases of evolutionary changes in the initial number of proliferating cells at growth plates. Future research on the genetic basis of process heterochronies and heteroposies will complete our understanding of these evolutionary phenomena.     

Combinatorial model for the formation of body plans in higher metazoan taxa: Paleontological insight

The body plans of higher metazoan taxa were formed during a short time (on the geological time scale) by combination of the previously developed characters. The combinations were realized as a result of manifestation of latent characters in adults through various heterochronies. This resulted in mosaic evolution and concealment of intermediate forms. Many characters of new body plans appeared in the ancestral taxon and their various combinations in the newly established taxa formed the archaic diversity. The maximum rank of newly appearing higher taxa decreased with geological time. The evolution of metazoans passed from the development of the general body plan to less significant details and appearance of body plans describing taxa of lower ranks. New body plans of higher taxa were superposed on the old body plan rather than replaced it, extending with time the subordination of body plans and respective hierarchy of taxa. Aromorphoses are always connected with the appearance of a new body plan. The appearance of new taxa and an increase in morphological diversity mostly occurred at certain boundaries in the development of the biota, which were connected with a sharp increase in the previously limited resources.     

Experimental approach to the hypotheses of heterochronic evolution in lower vertebrates

Heterochronies, temporal changes in ancestral ontogeny, are proposed to play the major role in microand macroevolutionary transformations of lower vertebrates. However, the evolutionary role of heterochronies often remains hypothetical, not verified experimentally. In the present paper, participation of heterochronies in (1) the origin of lacustrine fish species flocks, (2) the diversification of skeletal morphology in teleosts, and (3) the skull evolution in amphibians is experimentally verified. For this purpose, the temporal parameters of ontogeny were directly changed via artificial alterations of the thyroid hormones level in different representatives of lower vertebrates. The data obtained indicate that heterochronies are among the main mechanisms responsible for the current morphological diversity displayed by lower vertebrates at different phylogenetic levels.     

An integrative approach identifies developmental sequence heterochronies in freshwater basommatophoran snails

Adopting an integrative approach to the study of sequence heterochrony, we compared the timing of developmental events encompassing a mixture of developmental stages and functional traits in the embryos of 12 species of basommatophoran snails in an explicit phylogenetic framework. PARSIMOV analysis demonstrated clear functional heterochronies associated both with basal branches within the phylogeny and with terminal speciation events. A consensus of changes inferred under both accelerated transformation and delayed transformation optimizations identified four heterochronies where the direction of movement was known plus six twin heterochronies where the relative movements of the two events could not be assigned. On average, 0.5 and 0.58 events were inferred to have changed their position in the developmental sequence on internal and terminal branches of the phylogeny, respectively; these values are comparable with frequencies of sequence heterochrony reported in mammals. Directional heterochronies such as the early occurrence of body flexing in relation to the ontogeny of the eye spots, heart beat, and free swimming events occurred convergently and/or at different levels (i.e., familial, generic, and species) within the phylogeny. Such a functional approach to the study of developmental sequences has highlighted the possibility that heterochrony may have played a prominent role in the evolution of this group of invertebrates.     

Heterochronies,heterotopies, and cell resources of development in ontogenetic and evolutionary transformations

Classification and gene regulation of heterochronies and heterotopies as temporal and spatial evolutionary changes are considered. The intensity and duration of stem cell proliferation of a developing organism are connected with heterochronies and give rise to heterotopies. It is proposed that evolutionary emergence of new populations of stem cells (local embryomorphosis) is connected with heterotopies and heterochronies of gene expression, which support cell proliferation and temporarily suppress differentiation. The concept of modularity of evolutionary developmental biology and the theory of reproductive strategy naturally supplement each other and provide a better understanding of some ontogenetic changes in evolution. Developmental, morphological, and functional features of nematodes and vertebrates display alternative evolutionary trends with the predominance of progenesis and neoteny, respectively.     

MOLECULAR HETEROCHRONIES AND HETEROTOPIES IN EARLY ECHINOID DEVELOPMENT

By comparing the spatial and temporal distribution of three proteins during early development in seven echinoid species, we demonstrate that both heterochronies and heterotopies in gene-product expression have accompanied the radiation of post-Paleozoic echinoids. All three proteins examined showed significant alterations in time of expression, site of expression, or both. These molecular heterochronies and heterotopies indicate that early development is not necessarily as evolutionarily conservative as morphology of embryos alone would suggest. Evolutionary alterations in early development may be more common than is generally assumed.     

Modularity and heterochronies in the evolution of metazoa: Paleontological aspect

All organisms are formed of more or less independent elements, modules. Paleontology deals with morphological modules preserved in the fossil state and allows their evolution within taxa of different levels to be reconstructed. Modularity provides organisms with the ability to evolve, since changes in one module does not influence others, nor disturb the integrity of organism. Each organism may have unique modules represented by a single copy and serial modules developing according to a certain symmetry type. Serial terminal growth is characteristic of ambulacra of echinoderms, in which it is combined with alternating appearance of structures on the right and left of the symmetry plane. The morphology of the solute Maennilia estonica, which has been investigated in detail, shows that the growth model for the brachiola is similar to the model for ambulacra of sea urchins, but without an ocular plate. Probably, the hydrocoel initially induced the appearance of a skeleton necessary for its activity and organized its development according to its own model of terminal growth. Subsequently, the axial skeleton appearing following this pattern could have organized the growth of adjacent parts of the extraxial skeleton following the same model to form a united module. The fusion of modules could have resulted from heterochronies. Extant and extinct material connected with the change in the anteroposterior axis in evolutionary and ontogenetic development of echinoderms provides a prominent example of heterochronies. Heterochronies were the mechanism connecting characters into an integrated ensemble of the body plan. Archaic diversity reflects an attempt to create a new body plan. Various manifestations of archaic diversity show that the emergence of a new higher taxon is connected with the combination of a number of characters united in an integrated complex forming the body plan which is stable from the moment of appearance due to strict recursive relationships between its modules rather than with the acquisition of an individual character, even if it is very important.     

Methods for the analysis of developmental sequence data

SUMMARY Heterochrony, evolutionary changes in developmental timing, can be studied either by examining changes in growth or changes in the sequence of developmental events. Developmental sequence data has the potential to address many important questions in the field of developmental evolution, but methodological challenges remain due to the biological and logical dependence of events in a ranked sequence. In the past 10 years, the study of sequence heterochrony has undergone a rebirth, with the creation of several new methods for the analysis of this type of data. These methods can be divided into two broad categories: phenetic comparisons between terminal taxa that strive to uncover integrations within the developmental sequences and putative shared sequence heterochronies, and phylogeny-based methods that derive ancestor-descendent sequence heterochronies and establish statements of sequence evolution. In this review, we will discuss the strengths and weaknesses of the methodologies that have been proposed to quantitatively examine developmental sequence data, and studies that have attempted to implement them in an evolutionary context.     

Evolutionary mechanisms: Modularity,morphogenetic fields of gene expression,genetic regulation

Modern concepts heterochrony mechanisms, taking into account the data on modularity of ontogenetic and evolutionary processes, morphogenetic fields of gene expression are considered. In the context of evolutionary changes, features of genetic regulation of heterochronies, and also suppression of gene activity by epigenetic regulation are analyzed. Features of the origin of evolutionary novelties due to heterochronies, macromutations, and divergence of duplicated genes, which result in the formation of new genes and gene families, are discussed.     

Types of feeding in the ontogeny of fishes and the life-history model

Synopsis Different types of ontogenies in fishes-indirect and direct- are correlated with different nutrient availability and feeding during early life history. A comprehensive life-history model, developed earlier, facilitates the understanding of decisive events in the life of an organism. Embryos with insufficient endogenous food supply (yolk) to build a definitive phenotype directly need the transient form of a nutrient-gathering larva. They represent an indirect development. In contrast, a large endogenous supply of nutrients enables the definitive adult phenotype to develop directly, avoiding an intervening larva and the cost of metamorphosis. The larger and more advanced an individual at the onset of exogenous feeding, the better are its chances to survive. This can be achieved by heterochronies related to feeding. Different types of feeding during the early ontogeny of fishes — endogenous, exogenous, absorptive, and a combination of all (mixed) — are demonstrated and integrated into the life-history model.     

Influence of development rate on pharyngeal teeth formula in <Emphasis Type="Italic">Abramis brama</Emphasis> (L.) bream: Experimental cata

An experiment on acceleration and retardation of ontogeny with thyroid manipulation has revealed direct changes in definitive dentition of pharyngeal bones in Abramis brama bream. As development rate accelerates, the number of teeth reduces to the formula 5-4. When development rate is retarded, this number increases to the formula 6-5. Moreover, an additional minor row of teeth (1.6–5.1, 2.6–5.2) is formed. The observed changes in tooth numbers exceed the known variability in natural populations of bream. It is assumed that heterochronies lead to the changes in the number of teeth.     

Interpopulation comparison of variations in gastrulation of Rana arvalis and Rana ridibunda occurring under the conditions of anthropogenic contamination of water bodies

The results of morphometric analysis of gastrulation in city populations of Rana arvalis and Rana ridibunda suggest that under the conditions of anthropogenic contamination of water bodies, the early development is more variable, if we compare less variable sizes of their eggs in the city and suburb populations. The variability of gastrulation appears to depend on the degree of contamination of spawning water bodies and is accompanied by increased correlation between the forming structures. This latter may be determined by two causes: (1) increased number of interdependent features, which leads to coordinated differentiation of structures and regulation of variability, and (2) increased coefficients of correlation within a small group of features, which leads to the formation of structures characteristic for the subsequent developmental stages, including the appearance of heterochronies.     

Les relations ontogenèse-phylogenèse:Applications paléontologiques

After a short historical introduction, the principal notions used to study the connections between ontogeny and phylogeny are desribed. The three fondamental standards (shape, age, size) upon which are made ontogenetic comparisons are defined and discussed.The different ontogenetic heterochronies are described and illustrated with concrete paleontological examples. Successively are studied:u

1. -dwarfirsm and giantism which are particularcases;
2. -the 4 types of heterochrony which affect somatic development (neoteny and acceleration) or sexual maturation (hypermorphosis and progenesis);
3. -the innovations which are a fast appearence of a new character and their associations with the 4 elementary heterochronies.

The heterochronies are connected with phylogeny. This leads to the definition of two fondamental tendancies: paedomorphosis and peramorphosis; the terms proterogenesis and palingenesis are redefined. To conclude, this study is inserted in a more wider context that shows the importance of ontogenetic knowledge to understand numerous evolutif processes (tempos in evolution, demographic strategies, cladism).     

Deciphering the environmental impact on spike architectural traits for grain yield consolidation in bread wheat (T. aestivum L.)

The realization of grain yield in wheat is decided by source-sink balance under prevailing environmental conditions. Management conditions like changing the sowing time influence the source-sink capacity through modification in agronomic traits. Therefore, this experiment was conducted to decipher the influence of spike architectural traits (SATs) on grain yield and to open avenues for further grain yield enhancement. Comparatively early sowing over timely sowing gives the advantage of realizing higher grain yield with a positive relationship with SATs namely spike length, spikelets per spike, individual spike weight, individual grain weight, number of grains per spikelet, grain length, and grain width of upper and lower spike portion. Confirmatory factorial analysis revealed that spike length, spikelets per spike, individual spike weight, grains per spikelet were having a significant effect in deciding grain yield in early sown. The presence of a significant effect of genotype by environment interaction over grain yield and SATs allows the exploitation of available genotypic and environmental variability for further yield enhancement. GGE analysis on transformed and standardized grain yield-trait (GY-trait) combinations was used in the selection of genotypes having high GY-trait combinations for both sowing times. In early sowing, WG 11 was the best for high GY with high individual spike weight; grain length and grain width at lower and upper parts of the spike; and shorter days to 50% flowering. Genotypes exclusively having the high GY-trait combination along with low values of remaining GY-trait combinations were also selected with genotype focused GGE approach.     

Nearest-neighbor tree species combinations in tropical forest: the role of chance, and some consequences of high diversity

In three permanent inventory plots comprising 12.4 ha of undisturbed forest at La Selva, Costa Rica, all stems ≥10 cm dbh were mapped and identified to species. There were 1628, 1478 and 1954 trees in the plots, representing 168, 166 and 171 species respectively. We determined the species of each nearest-neighbor pair of trees, and asked whether the occurrence of species pairs conforms to a simple random mixing model. If trees are randomly mixed in terms of species, the expected frequency of any nearest neighbor species combination is a function of the relative abundance of the two species. Departures from random mixing could arise from species interactions, differential responses to habitat, or both. The number of possible ij species combinations increases approximately as the square of the number of species. For the 168 species in plot 1, for example, there are 14 196 possible combinations. We compared the expected frequency of each species combination in the three plots (42 736 combinations in all) with observed frequencies. Over 98% of the combinations had observed frequencies of zero and expected frequencies close to zero. A consequence of high diversity is low density of most individual species, and exceedingly low frequencies of the vast majority of species combinations. For each of the 805 combinations with observed frequencies >0, we used simulation to generate a distribution of expected frequencies. We used a t-test to compare the observed frequency with the mean of the simulated distribution for each combination. Only 40 combinations (0.09% of the possible species combinations in the plots) departed from expected frequencies; 39 combinations were more common, and one less common than expected. The overwhelming majority of nearest neighbor species combinations occur at frequencies predictable from their individual abundances.     

Short- and long-term advantages of an alternative ontogenetic pathway

We documented hypotheses on the evolution of developmental heterochronies by estimating short- and long-term advantages of alternative morphs. In this respect, we compared food energy intakes and body condition between paedomorphic and metamorphic Alpine newts ( Triturus alpestris , Caudata, Amphibia) in four populations. Because we found a strong correlation between fat reserves and body condition, we used this last parameter as an indicator of long-term gains. In all studied sites, paedomorphic females showed higher body condition than metamorphic ones. Paedomorphic males were also at advantage in two populations. Paedomorphs exhibited higher energy intakes than metamorphs in two populations and similar gains in two others. Our results support unifying theories that predict the occurrence of facultative paedomorphosis in varied habitats as paedomorphs exhibited better energy balance than metamorphs. This work shows the need for considering integrative parameters such as body condition in the comparative study of performances when studying developmental heterochronies. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 105–112     

Early embryogenesis in discoglossoid frogs: a study of heterochrony at different taxonomic levels

We studied the timing of the early development of the cranial neural crest, neural tube differentiation, somite formation and body elongation in the closely related frog species Discoglossus montalenti , Discoglossus galganoi , Discoglossus pictus , Bombina orientalis , Bombina variegata and Bombina maxima . Despite their different egg and embryo sizes, event pairing shows that there are only very few heterochronies among the three Bombina species and the three Discoglossus species in early developmental events. Instead, differences become apparent at the genus level (interpreted as phylogenetic signal). For example, the separation into a mandibular and a hyoid portion of cranial neural crest could be seen earlier in the Discoglossus than in the Bombina species when compared with the development of the neural tube. The development of the cranial neural crest streams also shifts (relatively) forward in time when compared with body elongation and somite formation in the Discoglossus species in contrast to the Bombina species. Comparisons with other species show that attempts to correlate heterochronies in early development with life history changes should take into account the possibility of a rather high evolvability, i.e. a high probability of convergence in the timing of these early events.     

Timing of ossification in duck, quail, and zebra finch: intraspecific variation, heterochronies, and life history evolution

Skeletogenic heterochronies have gained much attention in comparative developmental biology. The temporal appearance of mineralized individual bones in a species - the species ossification sequence - is an excellent marker in this kind of study. Several publications describe interspecific variation, but only very few detail intraspecific variation. In this study, we describe and analyze the temporal order of ossification of skeletal elements in the zebra finch, Taeniopygia guttata, the Japanese quail, Coturnix coturnix japonica, and the White Pekin duck, a domestic race of the mallard Anas platyrhynchos, and explore patterns of intraspecific variation in these events. The overall sequences were found to be conserved. In the duck, variability is present in the relative timing of ossification in the occipital, the basisphenoid and the otic regions of the skull and the phalanges in the postcranium. This variation appears generally in close temporal proximity. Comparison with previously published data shows differences in ossification sequence in the skull, the feet, and the pelvis in the duck, and especially the pelvis in the quail. This clearly documents variability among different breeds.     

小麦新品种皖麦17选育的理论与实践

本文对一个高产小麦新品种选育实践进行了剖析。结果表明：在Ｆ１按杂种的综合性状进行指数估算,然后把指数当作一个性状,估算各组合的杂种优势和配合力,并以此评判组合,可以增加对组合认识的预见性。再根据单株表现分别种植,可以在Ｆ３形成系统群。Ｆ２株系间则可表现出遗传差异,结合测产选择单株,并以对照单株综合指数为标准,再把高产组合作为选择（株行或单株）重点,可提高杂种的选择效率。Ｆ３以后一边鉴定株系,一边继续单株选择,可使品系纯化和提高同步深化。Ｆ６以后继续优中选优仍然能使增效基因积累。