On Novelty,Heterochrony and Developmental Constraints in a Complex Morphological Theory of Recapitulation: The Genus <Emphasis Type="Italic">Trophon</Emphasis> as a Case Study

Darwin proposed natural selection as the main evolutionary mechanism in 1859. However, he did not think that this was the only process by which new species were generated. It was the so-called Modern Synthesis who established natural selection as the only mechanism responsible for evolution. Since then, the evolutionary process is explained by the pair mutation-adaptation: new species are generated by the appearance of new mutations, which in case of allowing new adaptations to the environment, they will be fixed and organisms will survive, therefore resulting in new species. An alternative view to the plasticity promoted by the adaptationist program is to think organisms as truly organized structures, having different levels of structural organization, which would mean that not every form is possible, but only those that correspond to a certain building plan. This would be reflected in the appearance of structural constraints, showing the limits imposed to the organism during its evolutionary development. In this work, I studied the ontogeny and development of three species of the genus Trophon by geometric morphometrics, in order to clarify important concepts in evolutionary developmental biology (Evo-Devo). Integrating theoretical and empirical investigations, I could propose a new conceptual framework for heterochrony in a context of a complex theory of recapitulation. Furthermore, I could detect a developmental constraint in Trophon, which provided an opportunity to reconstruct the concept of constraint and propose a synthesis between heterochrony and constraint that explained evolution as a process fueled by them, that is, as directive and driving force.     

The developmental renaissance in adaptationism

From an adaptation perspective, unoccupied patches of morphological space are inferred to be empty because they are of low fitness and selected against. These inferences hinge on venturesome assumptions, because emptiness is explained by low fitness and low fitness is inferred from emptiness. Moreover, non-adaptive factors, such as developmental constraint, could also plausibly account for empty morphospace. In response, biologists increasingly study ontogeny to test the assumption that unobserved phenotypes could be produced if selection were to favor them; finding that empty space morphologies can be readily produced in development helps reject constraint and lends support to adaptive hypotheses. This developmental approach to adaptation calls on manifold techniques, including embryology, artificial selection and comparative methods. Belying their diversity, all of these methods examine the causes of empty morphospace and mark a return of development, long excluded from traditional evolutionary biology, to adaptationist practice.     

Evolutionary implications of developmental instability in parthenogenetic drosophila mercatorum. I. Comparison of several strains with different genotypes

Natural populations of sexually reproducing Drosophila mercatorum are capable of a very low rate of parthenogenesis, but this mode of reproduction has apparently never characterized an entirely asexual population in this species. The high abortion rate observed in laboratory parthenogenetic lines suggests that developmental constraints may cause the failure of this trait to spread in nature. To investigate the basis of this developmental instability and how it may affect the evolution of parthenogenesis in natural populations, early embryonic development was compared between one sexual and four parthenogenetic laboratory strains of D. mercatorum. There is a large amount of variation within a given parthenogenetic strain, suggesting that parthenogenesis is associated with a general breakdown of developmental stability. There is relatively little variation among different parthenogenetic strains, suggesting that most abortions are due to a feature inherent to parthenogenetic reproduction rather than a feature of a particular genome. Likewise, there is little variation between parthenogenetic and sexual strains in the causes of abortions, suggesting that the developmental problems encountered by parthenogenetic lineages are not unique to parthenogens. Thus, the failure of parthenogenesis to spread within D. mercatorum can be attributed to no particular developmental constraint per se operating after the initiation of embryogenesis. However, the overall increase in all developmental problems that occurs with the transition from sexual to parthenogenetic development suggests that the high degree of developmental instability associated with parthenogenesis may be considered a developmental constraint in its own right.     

Matters of scale: positive allometry and the evolution of male dimorphisms

The developmental independence of alternative phenotypes is key to evolutionary theories of phenotypic plasticity and the origins of diversity. Male dimorphisms associated with alternative reproductive tactics are widely cited examples of such facultative expression of divergent fitness optima. Current models for the evolution of male dimorphisms invoke a size-dependent threshold at which the phenotype is reprogrammed. We use predictions derived from allometric modeling to test for the existence of reprogramming thresholds in two species of beetle, Onthophagus taurus and Onthophagus binodis, and the European earwig Forficula auricularia. We also compare the allometry of a number of morphological traits to determine whether minor males suppress their secondary sexual traits. The intercept of the horn allometry was suppressed, but there was no evidence of reprogramming of horn growth in either beetle species. There was reprogramming in the earwig. In the beetles, the horn length in all males can be explained largely in terms of exponential horn growth following an extraordinarily steep power function. The asymptote in O. taurus can be explained by exponential growth meeting the constraint of resource exhaustion. These findings question the currently held view that beetle horn dimorphisms showcase the importance of developmental independence in the evolution of diversity.     

Plant Succession and Tree Architecture: An Attempt at Reconciling Two Scales of Analysis of Vegetation Dynamics

Plant succession is a phenomenon ascribed to vegetation dynamics at the scale of the plant community. The study of plant succession implies the analysis of the species involved and their relationships. Depending on the research done, the characteristics of trees have been studied according to either static, dimensional or partial approaches. We have revised the principal theories of succession, the methods of describing structure and development of tree and relationship established between tree species' attributes and their successional status. During studies of succession, we propose that the profile of plant species be completed by including the organizational attributes of their structure as well as their development dynamics. We suggest an approach to the architecture that gives a general overview of the mode of development, revealing, among other things, that a limited number of architectural models may be found in nature. We hypothesize that community dynamics, in succession, may to a certain degree arise from the dynamic nature of plant development. This line of reasoning is in accordance with the current school of thought on process structuralism.     

Developmental constraints in cave beetles

In insects, whilst variations in life cycles are common, the basic patterns typical for particular groups remain generally conserved. One of the more extreme modifications is found in some subterranean beetles of the tribe Leptodirini, in which the number of larval instars is reduced from the ancestral three to two and ultimately one, which is not active and does not feed. We analysed all available data on the duration and size of the different developmental stages and compared them in a phylogenetic context. The total duration of development was found to be strongly conserved, irrespective of geographical location, habitat type, number of instars and feeding behaviour of the larvae, with a single alteration of the developmental pattern in a clade of cave species in southeast France. We also found a strong correlation of the size of the first instar larva with adult size, again regardless of geographical location, ecology and type of life cycle. Both results suggest the presence of deeply conserved constraints in the timing and energy requirements of larval development. Past focus on more apparent changes, such as the number of larval instars, may mask more deeply conserved ontogenetic patterns in developmental timing.     

Germination and regeneration mechanisms in Mediterranean degenerate forests

Abstract. Germination and developmental behaviour during juvenile stages are of importance in relation to succession mechanisms acting on plant communities. This is particularly so in the Mediterranean region, where the climatic summer drought causes stress. We investigated and compared germination and development of four tree species in two parts of the French Mediterranean region where the original dominant tree species (Quercus pubescens) has been largely replaced by Quercus ilex in the southern part, and Castanea sativa in the northern part. Observed differences in germination rates and developmental behaviour, especially shootroot ratio and the role of water supply, suggest that other species may eventually replace the current dominant species.     

Secondary succession and summer herbivory in a subarctic grassland: community structure and diversity

A field experiment was established in a subarctic grassland in the Finnish Lapland to study the role of summer herbivory in plant community succession Perennial vegetation and moss cover were removed in an area of 324 m² The site was divided into four blocks, of which two were fenced to prevent herbivory by large mammals (reindeer, hare)
Early successional changes in the vegetation were assessed Mean species richness per 3 × 3 m plot was consistently higher in the fenced area, indicating that herbivory can suppress small-scale diversity Herbivory affected the height of several plant species However, there was no correlation between frequency and height of individual species There was a weak indication that taller species were more successful m early succession when grazed Light competition is apparently not a key process determining successional change Thus, in early stage of succession, summer herbivory has little effect on diversity by limiting light competition, and most species are equally successful in grazed and ungrazed plots There was some indirect evidence about competitive interactions in the developing community However, unlike temperate grasslands, large mammal herbivory and competition for light seem not to be important determinants of community change in this subarctic grassland (at least what concernes early successional stages) This may be explained by the harshness of local climate, and abundance of light due to the polar day     

Relations between Phytoplankton and Light Climate in two Swedish Forest Lakes

The succession, biomass and production of phytoplankton was studied during 7 years in two Swedish forest lakes in relation to nutrients, temperature and light climate. The succession of species and taxonomic groups can be described by their different demands concerning temperature and light climate while the total biomass is correlated to light climate and total phosphorus concentration. Differences in production between the two lakes can be explained by different taxonomic groups having different efficiencies.     

Developmental constraints and convergent evolution in Drosophila sex comb formation

SUMMARY The most complex and diverse secondary sexual character in Drosophila is the sex comb (SC), an arrangement of modified bristles on the forelegs of a subclade of male fruit flies. We examined SC formation in six representative nonmodel fruit fly species, in an effort to understand how the variation in comb patterning arises. We first compared SC development in two species with relatively small combs, Drosophila takahashii , where the SCs remain approximately transverse, and Drosophila biarmipes , where two rows of SC teeth rotate and move in an anterior direction relative to other bristle landmarks. We then analyzed comb ontogeny in species with prominent extended SCs parallel to the proximodistal axis, including Drosophila ficusphila and species of the montium subgroup. Our study allowed us to identify two general methods of generating longitudinal combs on the tarsus, and we showed that a montium subgroup species ( Drosophila nikananu ) with a comb convergently similar in size, orientation and position to the model organism Drosophila melanogaster , forms its SC through a different developmental mechanism. We also found that the protein product of the leg patterning gene, dachshund (dac) , is strongly reduced in the SC in all species, but not in other bristles. Our results suggest that an apparent constraint on SC position in the adult may be attributable to at least two different lineage-specific developmental processes, although external forces could also play a role.     

Do precocial mammals develop at a faster rate? A comparison of rates of skull development in Sigmodon fulviventer and Mus musculus domesticus

Variation in neonatal maturity among mammals is often explained by variation in gestation length, but species may also differ in developmental rate, a quantity that is difficult to measure because the conventional formalism makes two important and potentially unrealistic assumptions: (1) ontogeny of form can be described by a single line, and (2) species have the same ontogeny of form. We examine two species, one precocial (Sigmodon fulviventer), the other altricial (Mus musculus domesticus), and find that neither assumption is met. Therefore, we introduce an alternative metric, the rate of shape differentiation away from the average neonate. We find that S. fulviventer has a lower developmental rate than M. m. domesticus; consequently, while more mature at birth, S. fulviventer loses ground to M. m. domesticus over time. Surprisingly, despite differences in gestation length and developmental rate, these species reach developmental and life-history milestones at nearly identical degrees of skull shape maturity.     

羊草草地放牧退化演替中种群消长模型的研究

羊草草地是东北西部及内蒙东部草原区主要的天然放牧场和割草场、具有较高的经济价值,但是由于长期过度放牧,草地已严重退化并盐碱化。本文采用数学手段,以植物种群在群落中的相对优势度为指标,用植被在放牧退化演替系列上的空间分布序列推断其在时间上演替系列的方法,研究羊草等5种植物种群的优势度随放牧退化演替消长的规律及其数学模型。羊草按D＝124.2e-0.35T衰退,寸草苔和糙隐子草分别按D＝2.13T3.3e-0.15T2和 D=1.77T3.7e-0.29T2消长,而角碱蓬和虎尾草分别按D=0.11e1.3T和D=0.38e0.94T增加。     

Parthenogenesis and developmental constraints

The absence of a paternal contribution in an unfertilized ovum presents two developmental constraints against the evolution of parthenogenesis. We discuss the constraint caused by the absence of a centrosome and the one caused by the missing set of chromosomes and how they have been broken in specific taxa. They are examples of only a few well‐underpinned examples of developmental constraints acting at macro‐evolutionary scales in animals. Breaking of the constraint of the missing chromosomes is the best understood and generally involves rare occasions of drastic changes of meiosis. These drastic changes can be best explained by having been induced, or at least facilitated, by sudden cytological events (e.g., repeated rounds of hybridization, endosymbiont infections, and contagious infections). Once the genetic and developmental machinery is in place for regular or obligate parthenogenesis, shifts to other types of parthenogenesis can apparently rather easily evolve, for example, from facultative to obligate parthenogenesis, or from pseudoarrhenotoky to haplodiploidy. We argue that the combination of the two developmental constraints forms a near‐absolute barrier against the gradual evolution from sporadic to obligate or regular facultative parthenogenesis, which can probably explain why the occurrence of the highly advantageous mode of regular facultative parthenogenesis is so rare and entirely absent in vertebrates.     

Evolutionary origins of novel conchologic growth patterns in tropical American corbulid bivalves

We conducted a combined sclerochronologic and phylogenetic analysis to document patterns and rates of shell accretion in several subclades of related corbulids, and to explore the evolutionary origins of novel conchologic developmental patterns. We found three disparate patterns of valve development in Neogene tropical American corbulid genera. These patterns include growth through primarily radial accretion along the sagittal plane, and two derivative patterns: one characterized by initial deposition of a thin shell followed by valve thickening with little increase in valve height, and another producing a well-defined nepioconch through a marked change in the primary growth direction. We conducted a species-level phylogenetic analysis of the taxa surveyed for growth patterns, focusing on the ([Bothrocorbula+Hexacorbula]+Caryocorbula) clade. The phylogenetic distribution of shell growth patterns suggests that this clade is characterized by derivative patterns of growth. Oxygen-isotope calibrated ontogenetic age estimates of species in the derived Bothrocorbula subclade further suggest that transitions from the ancestral radial (sagittal) growth pattern to a derived pattern of growth are a function of heterochrony (peramorphosis by acceleration). These findings are significant because they link previously observed patterns of morphological constraint with a specific evolutionary process, demonstrate how morphologic constraint and innovation can be interrelated, and serve as a model for understanding the evolution of morphologic diversity in the clade as a whole. Furthermore, this study highlights the utility of sclerochronologic records as an important component of evolutionary developmental research on organisms with accretionary skeletal growth.     

Primitive versus derived traits in the developmental program of the vertebrate head: views from cyclostome developmental studies

Evolution can be viewed as a series of changes in the developmental program along the phylogenetic tree. To better understand the early evolution of the vertebrate skull, we can use the embryos of the cyclostome species as models. By comparing the cyclostome developmental patterns with those of gnathostomes, it becomes possible to distinguish the primitive and derived parts of the developmental program as taxon-specific traits. These traits are often recognizable as developmental constraints that define taxa by biasing the developmental trajectories within a certain limited range, resulting in morphological homologies in adults. These developmental constraints are distributed on the phylogenetic tree like the morphological character states of adult animals and are associated with specific regions of the tree. From this perspective, we emphasize the importance of considering gene expression and embryonic anatomy as the mechanistic bases that can result in homologous or nonhomologous morphological patterns at later developmental stages. Taking the acquisition of the jaw and trabecula cranii as examples, we demonstrate that a set of embryonic features can be coupled or decoupled during evolution and development. When they are coupled, they exert an ancestral developmental constraint that results in homologous morphological patterns, and when they are decoupled, the ancestral constraints tend to be abandoned, generating a new body plan. The heterotopy behind the specification of the oral domain is an example of decoupling, based on shifted tissue interactions. We also stress the importance of "developmental burden" in determining the sequential order of changes through evolution.     

Stochastic trajectories of succession initiated by extreme climatic events

Deterministic or rule-based succession is expected under homogeneous biotic and abiotic starting conditions. Effects of extreme climatic events such as drought, however, may alter these assembly rules by adding stochastic elements. We monitored the succession of species composition of 30 twin grassland communities with identical biotic and abiotic starting conditions in an initially sown diversity gradient between 1 and 16 species over 13 years. The stochasticity of succession, measured as the synchrony in the development of the species compositions of the twin plots, was strongly altered by the extreme warm and dry summer of 2003. Moreover, it was independent from past and present plant diversity and neighbourhood species compositions. Extreme climatic events can induce stochastic effects in community development and therefore impair predictability even under homogeneous abiotic conditions. Stochastic events may result in lasting shifts of community composition, as well as adverse and unforeseeable effects on the stability of ecological services.     

Phenoptosis in arthropods and immortality of social insects

In general, there are no drastic differences in phenoptosis patterns in plant and animal organisms. However, there are some specific features characteristic for insects and other arthropods: 1) their development includes metamorphosis with different biochemical laws at consecutive developmental stages; 2) arthropods can reduce or stop development and aging when in a state of diapause or temporal cold immobility; 3) their life cycle often correlates with seasonal changes of surroundings; 4) polymorphism is widespread — conspecifics differ by their lifespans and phenoptosis features; 5) lifespan-related sexual dimorphism is common; 6) significant situational plasticity of life cycle organization is an important feature; for example, the German wasp (Paravespula germanica) is obligatorily univoltine in the temperate zone, while in tropical regions its lifespan increases and leads to repeated reproduction; 7) life cycles of closely related species may differ significantly, for example, in contrast to German wasp, some tropical hornets (Vespa) have only one reproduction period. Surprisingly, many insect species have been shown to be subjected to gradual aging and phenoptosis, like the highest mammals. However, queens of social insects and some long-lived arachnids can apparently be considered non-aging organisms. In some species, lifespan is limited to one season, while others live much longer or shorter. Cases of one-time reproduction are rather rare. Aphagia is common in insects (over 10,000 species). Cannibalism is an important mortality factor in insects as well as in spiders. In social insects, which exist only in colonies (families), the lifetime of a colony can be virtually unlimited. However, in case of some species the developmental cycle and death of a colony after its completion are predetermined. Most likely, natural selection in insects does not lengthen individual lifespan, but favors increase in reproduction efficiency based on fast succession of generations leading to increased evolvability.     

Developmental drive: an important determinant of the direction of phenotypic evolution

SUMMARY Over any period of evolutionary time, the prevailing ontogenetic trajectory within a lineage may either recur unchanged from generation to generation (stasis) or alter (developmental reprogramming). A key question about reprogramming is whether it exhibits intrinsic biases in favor of some sorts of change and against others, which may be referred to respectively as "drive" and "constraint." A simple logical argument suggests that both drive and constraint should be common, and conversely that cases of equiprobable modification in various phenotypic directions should be relatively rare. These proposals, that drive and constraint exist and that they are common, appear to be widely accepted, even among neo-Darwinians, who are sometimes portrayed as rejecting them. What is more controversial is that developmental drive (and constraint) can have a powerful influence on the direction of evolutionary change. It is argued that such an influence will occur, and indeed may be pervasive.     

A DEVELOPMENTAL ANALYSIS OF AN EVOLUTIONARY TREND: DIGITAL REDUCTION IN AMPHIBIANS

In this study, we integrate information from phylogeny, comparative ontogeny, and experimental embryology in an attempt to elucidate the mechanisms controlling evolutionary trends towards digital reduction and loss observed in amphibians. Frogs and salamanders that have lost phalanges and even whole toes have done so in a very ordered manner, i.e., certain skeletal elements are lost prior to others. This pattern of morphological diversity is described and trends elucidated. It is concluded that the process is characterized by striking intraordinal convergences coupled with substantial differences between the trends observed in frogs as compared to urodeles. We argue that this pattern is essentially a reflection of the differences in the ontogenies of the two orders. Similarly, the convergences within urodeles and within anurans can be explained as the result of regulation of developmental parameters in a resilient developmental program. We further explore this hypothesis by experimentally perturbing the number of cells in the embryonic limb primordium to show that reduction in the number of mesenchymal cells secondarily affects the developmental process of pattern formation causing a rearrangement of the skeletal morphology of the foot. The same experimental manipulation has different effects in frogs as compared to salamanders. However, in both cases, the experimentally generated morphologies tend to parallel the phenotypes and trends observed in nature. Our conclusion is that most of the patterns of diversity in the digital morphology of amphibians can be explained as a reflection of developmental properties. In general, we present a methodology that attempts to empirically address the issue of identifying developmental constraint in morphological evolution.