The origins and the early evolution of multicellular animals required the exploitation of holozoan genomic regulatory elements and the acquisition of new regulatory tools. Comparative studies of ...metazoans and their relatives now allow reconstruction of the evolution of the metazoan regulatory genome, but the deep conservation of many genes has led to varied hypotheses about the morphology of early animals and the extent of developmental co-option. In this Review, I assess the emerging view that the early diversification of animals involved small organisms with diverse cell types, but largely lacking complex developmental patterning, which evolved independently in different bilaterian clades during the Cambrian Explosion.
The history of life as documented by the fossil record encompasses evolutionary diversifications at scales ranging from the Ediacaran–Cambrian explosion of animal life and the invasion of land by ...vascular plants, insects and vertebrates to the diversification of flowering plants over the past 100 million years and the radiation of horses. Morphological novelty and innovation has been a recurrent theme. The architects of the modern synthesis of evolutionary theory made three claims about evolutionary novelty and innovation: first, that all diversifications in the history of life represent adaptive radiations; second, that adaptive radiations are driven principally by ecological opportunity rather than by the supply of new morphological novelties, thus the primary questions about novelty and innovation focus on their ecological and evolutionary success; and third, that the rate of morphological divergence between taxa was more rapid early in the history of a clade but slowed over time as ecological opportunities declined. These claims have strongly influenced subsequent generations of evolutionary biologists, yet over the past two decades each has been challenged by data from the fossil record, by the results of comparative phylogenetic analyses and through insights from evolutionary developmental biology. Consequently a broader view of novelty and innovation is required. An outstanding issue for future work is identifying the circumstances associated with different styles of diversification and whether their frequency has changed through the history of life.
Douglas Erwin reviews the patterns of evolutionary diversifications, arguing that there is a wider array of patterns than predicted by models of adaptive radiation, and that the origins of evolutionary novelty may be independent of ecological opportunity.
ABSTRACT
Since 1990 the recognition of deep homologies among metazoan developmental processes and the spread of more mechanistic approaches to developmental biology have led to a resurgence of ...interest in evolutionary novelty and innovation. Other evolutionary biologists have proposed central roles for behaviour and phenotypic plasticity in generating the conditions for the construction of novel morphologies, or invoked the accessibility of new regions of vast sequence spaces. These approaches contrast with more traditional emphasis on the exploitation of ecological opportunities as the primary source of novelty. This definitional cornucopia reflects differing stress placed on three attributes of novelties: their radical nature, the generation of new taxa, and ecological and evolutionary impact. Such different emphasis has led to conflating four distinct issues: the origin of novel attributes (genes, developmental processes, phenotypic characters), new functions, higher clades and the ecological impact of new structures and functions. Here I distinguish novelty (the origin of new characters, deep character transformations, or new combinations) from innovation, the ecological and evolutionary success of clades. Evidence from the fossil record of macroevolutionary lags between the origin of a novelty and its ecological success demonstrates that novelty may be decoupled from innovation, and only definitions of novelty based on radicality (rather than generativity or consequentiality) can be assessed without reference to the subsequent history of the clade to which a novelty belongs. These considerations suggest a conceptual framework for novelty and innovation, involving: (i) generation of the potential for novelty; (ii) the formation of novel attributes; (iii) refinement of novelties through adaptation; (iv) exploitation of novelties by a clade, which may coincide with a new round of ecological or environmental potentiation; followed by (v) the establishment of innovations through ecological processes. This framework recognizes that there is little empirical support for either the dominance of ecological opportunity, nor abrupt discontinuities (often caricatured as ‘hopeful monsters’). This general framework may be extended to aspects of cultural and social innovation.
Recent molecular clock studies date the origin of Metazoa to 750–800 million years ago (Ma), roughly coinciding with evidence from geochemical proxies that oxygen levels rose from less than 0.1% ...present atmospheric level (PAL) to perhaps 1–3% PAL O2. A younger origin of Metazoa would require greatly increased substitution rates across many clades and many genes; while not impossible, this is less parsimonious. Yet the first fossil evidence for metazoans (the Doushantuo embryos) about 600 Ma is followed by the Ediacaran fossils after 580 Ma, the earliest undisputed bilaterians at 555 Ma, and an increase in the size and morphologic complexity of bilaterians around 542 Ma. This temporal framework suggests a missing 150–200 Myr of early metazoan history that encompasses many apparent novelties in the early evolution of the nervous system. This span includes two major glaciations, and complex marine geochemical changes including major changes in redox and other environmental changes. One possible resolution is that animals of these still unknown Cryogenian and early Ediacaran ecosystems were relatively simple, with highly conserved developmental genes involved in cell-type specification and simple patterning. In this model, complex nervous systems are a convergent phenomenon in bilaterian clades which occurred close to the time that larger metazoans appeared in the fossil record.
ABSTRACT
Environmental fluctuations in redox may reinforce rather than hinder evolutionary transitions, such that variability in near‐surface oceanic oxygenation can promote morphological evolution ...and novelty. Modern, low‐oxygen regions are heterogeneous and dynamic habitats that support low diversity and are inhabited by opportunistic and non‐skeletal metazoans. We note that several major radiation episodes follow protracted or repeating intervals (>1 million years) of persistent and dynamic shallow marine redox (oceanic anoxic events). These are also often associated with short‐lived mass‐extinction events (<0.5 million years) where skeletal benthic incumbents are removed, and surviving or newly evolved benthos initially inhabit transient oxic habitats. We argue that such intervals create critical opportunities for the generation of evolutionary novelty, followed by innovation and diversification.
We develop a general model for redox controls on the distribution and structure of the shallow marine benthos in a dominantly anoxic world, and compile data from the terminal Ediacaran–mid‐Cambrian (∼560–509 Ma), late Cambrian–Ordovician (∼500–445 Ma), and Permo‐Triassic (∼255–205 Ma) to test these predictions. Assembly of phylogenetic data shows that prolonged and widespread anoxic intervals indeed promoted morphological novelty in soft‐bodied benthos, providing the ancestral stock for subsequently skeletonized lineages to appear as innovations once oxic conditions became widespread and stable, in turn promoting major evolutionary diversification. As a result, we propose that so‐called ‘recovery’ intervals after mass extinctions might be better considered as ‘innovation’ intervals.
The link between biodiversity and climate has been obvious to biologists since the work of von Humboldt in the early 1800s, but establishing the relationship of climate to ecological and evolutionary ...patterns is more difficult. On evolutionary timescales, climate can affect supply of energy by biotic and abiotic effects. Some of the best evidence for a link between biodiversity and climate comes from latitudinal gradients in diversity, which provide an avenue to explore the more general relationship between climate and evolution. Among the wide range of biotic hypotheses, those with the greatest empirical support indicate that warmer climates have provided the energetic foundation for increased biodiversity by fostering greater population size and thus increased extinction resistance; have increased metabolic scope; have allowed more species to exploit specialized niches as a result of greater available energy; and generated faster speciation and/or lower extinction rates. In combination with geologic evidence for carbon dioxide levels and changing areas of tropical seas, these observations provide the basis for a simple, first-order model of the relationship between climate through the Phanerozoic and evolutionary patterns and diversity. Such a model suggests that we should expect greatest marine diversity during globally warm intervals with dispersed continents, broad shelves and moderately extensive continental seas. Demonstrating a significant evolutionary response to either climate or climatic change is challenging, however, because of continuing uncertainties over patterns of Phanerozoic marine diversity and the variety of factors beyond climate that influence evolution.
Comparative developmental evidence indicates that reorganizations in developmental gene regulatory networks (GRNs) underlie evolutionary changes in animal morphology, including body plans. We argue ...here that the nature of the evolutionary alterations that arise from regulatory changes depends on the hierarchical position of the change within a GRN. This concept cannot be accomodated by microevolutionary nor macroevolutionary theory. It will soon be possible to investigate these ideas experimentally, by assessing the effects of GRN changes on morphological evolution.
Celotno besedilo
Dostopno za:
DOBA, IJS, IZUM, KILJ, NUK, PILJ, PNG, SAZU, UILJ, UKNU, UL, UM, UPUK
Sewall Wright's fitness landscape introduced the concept of evolutionary spaces in 1932. George Gaylord Simpson modified this to an adaptive, phenotypic landscape in 1944 and since then evolutionary ...spaces have played an important role in evolutionary theory through fitness and adaptive landscapes, phenotypic and functional trait spaces, morphospaces and related concepts. Although the topology of such spaces is highly variable, from locally Euclidean to pre-topological, evolutionary change has often been interpreted as a search through a pre-existing space of possibilities, with novelty arising by accessing previously inaccessible or difficult to reach regions of a space. Here I discuss the nature of evolutionary novelty and innovation within the context of evolutionary spaces, and argue that the primacy of search as a conceptual metaphor ignores the generation of new spaces as well as other changes that have played important evolutionary roles.
This article is part of the themed issue ‘Process and pattern in innovations from cells to societies’.
That the activities of organisms influence their surrounding ecological communities, and the environment, has long been appreciated by palaeontologists, as has the role of these activities on both ...ecological and evolutionary processes. Spillover effects extend the range of ecosystem‐engineering through ecological networks, generating network effects that because of their non‐trophic nature can be challenging to track. Moreover, the cumulative effect of organismal activities can persist far beyond the lifespan of individual organisms, producing ecological inheritances that influence macroecological and macroevolutionary dynamics. This contribution surveys macroevolutionary patterns arising from ecosystem engineering, their potential contribution to evolutionary radiations, and the significance of ecosystem engineering as a public good in the success of evolutionary innovations. Anecdotally, such activities appear to have made important contributions, but considerable work is required for more rigorous understanding. I describe two challenges: the need for palaeontologists to collect abundance data in a way that facilitates comparative study, and the importance of more robust models of ecological (not just trophic) networks involving multigraphs and hypergraphs.
Diverse bilatería clades emerged apparently within a few million years during the early Cambrian, and various environmental, developmental, and ecological causes have been proposed to explain this ...abrupt appearance. A compilation of the patterns of fossil and molecular diversification, comparative developmental data, and information on ecological feeding strategies indicate that the major animal clades diverged many tens of millions of years before their first appearance in the fossil record, demonstrating a macroevolutionary lag between the establishment of their developmental toolkits during the Cryogenian (850 to 635 million years ago Ma) and the later ecological success of metazoans during the Ediacaran (635 to 541 Ma) and Cambrian (541 to 488 Ma) periods. We argue that this diversification involved new forms of developmental regulation, as well as innovations in networks of ecological interaction within the context of permissive environmental circumstances.
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