The innovation triad: an EvoDevo agenda

This article introduces a special issue on evolutionary innovation and morphological novelty, two interrelated themes that have received a remarkable increase of attention over the past few years. We begin with a discussion of the question of whether innovation and novelty represent distinct evolutionary problems that require a distinct conceptualization. We argue that the mechanisms of innovation and their phenotypic results--novelty--can only be properly addressed if they are distinguished from the standard evolutionary themes of variation and adaptation, and we present arguments for making such a distinction. We propose that origination, the first formation of biological structures, is another distinct problem of morphological evolution, and that together with innovation and novelty it constitutes a conceptual complex we call the innovation triad. We define a problem agenda of the triad, which separates the analysis of the initiating conditions from the mechanistic realization of innovation, and we discuss the theoretical problems that arise from treating innovation as distinct from variation. Further, we categorize the empirical approaches that address themes of the innovation triad in recognizing four major strands of research: the morphology and systematics program, the gene regulation program, the epigenetic program, and the theoretical biology program. We provide examples of each program, giving priority to contributions in the present issue. In conclusion, we observe that the innovation triad is one of the defining topics of EvoDevo research and may represent its most pertinent contribution to evolutionary theory. We point out that an inclusion of developmental systems properties into evolutionary theory represents a shift of explanatory emphasis from the external factors of natural selection to the internal dynamics of developmental systems, complementing adaptation with emergence, and contingency with inherency.     

EvoDevo and niche construction: building bridges

Evolutionary developmental biology and niche-construction theory have much in common, despite independent intellectual origins. Both place emphasis on the role of ontogenetic processes in evolution. The same historical events shaped them, and similar philosophical and sociological barriers hindered their respective advances. Both perspectives maintain that neo-Darwinism needs a theory of macroevolutionary variation and that such a theory can now be adduced from developmental biology. Some proponents of both EvoDevo and niche construction propose additional evolutionary mechanisms, and specify a key role for stable extra-genetic forms of inheritance. Similarly, proponents of each lay emphasis on "reciprocal causation" in the relationship between organism and environment. We illustrate here how EvoDevo and niche construction could gain "added value" from each other, and demonstrate how the niche-construction perspective potentially provides a useful conduit to integrate evolutionary and developmental biology.     

Phenotypic Novelty in EvoDevo: The Distinction Between Continuous and Discontinuous Variation and Its Importance in Evolutionary Theory

The introduction of novel phenotypic structures is one of the most significant aspects of organismal evolution. Yet the concept of evolutionary novelty is used with drastically different connotations in various fields of research, and debate exists about whether novelties represent features that are distinct from standard forms of phenotypic variation. This article contrasts four separate uses for novelty in genetics, population genetics, morphology, and behavioral science, before establishing how novelties are used in evolutionary developmental biology (EvoDevo). In particular, it is detailed how an EvoDevo-specific research approach to novelty produces insight distinct from other fields, gives the concept explanatory power with predictive capacities, and brings new consequences to evolutionary theory. This includes the outlining of research strategies that draw attention to productive areas of inquiry, such as threshold dynamics in development. It is argued that an EvoDevo-based approach to novelty is inherently mechanistic, treats the phenotype as an agent with generative potential, and prompts a distinction between continuous and discontinuous variation in evolutionary theory.     

人体微生态系统与人类宏基因组计划

介绍了微生态系统的基本概念和人体基本的微生态系统,阐述了其对人类的意义;介绍了人类宏基因组计划基本知识和研究方法.     

Delineation of Interfaces on Human Alpha-Defensins Critical for Human Adenovirus and Human Papillomavirus Inhibition

Human α-defensins are potent anti-microbial peptides with the ability to neutralize bacterial and viral targets. Single alanine mutagenesis has been used to identify determinants of anti-bacterial activity and binding to bacterial proteins such as anthrax lethal factor. Similar analyses of α-defensin interactions with non-enveloped viruses are limited. We used a comprehensive set of human α-defensin 5 (HD5) and human neutrophil peptide 1 (HNP1) alanine scan mutants in a combination of binding and neutralization assays with human adenovirus (AdV) and human papillomavirus (HPV). We have identified a core of critical hydrophobic residues that are common determinants for all of the virus-defensin interactions that were analyzed, while specificity in viral recognition is conferred by specific surface-exposed charged residues. The hydrophobic residues serve multiple roles in maintaining the tertiary and quaternary structure of the defensins as well as forming an interface for virus binding. Many of the important solvent-exposed residues of HD5 group together to form a critical surface. However, a single discrete binding face was not identified for HNP1. In lieu of whole AdV, we used a recombinant capsid subunit comprised of penton base and fiber in quantitative binding studies and determined that the anti-viral potency of HD5 was a function of stoichiometry rather than affinity. Our studies support a mechanism in which α-defensins depend on hydrophobic and charge-charge interactions to bind at high copy number to these non-enveloped viruses to neutralize infection and provide insight into properties that guide α-defensin anti-viral activity.     

Human teratocarcinomas

Teratocarcinomas are one of the commonest forms of cancer in young adult men. Cell lines derived from these tumors, and particularly the cell lines composed of their embryonal carcinoma (EC) stem cells, may provide useful information concerning the development and subsequent pathology of teratocarcinomas in humans. In addition, it is likely that human EC cells resemble early embryonic cells and can be used as an in vitro counterpart of such cells from the human embryo. Several common properties of human EC cells have been identified, and a human EC cell line, TERA-2, that is capable of extensive somatic differentiation has been cloned. In nude mice, TERA-2 EC cells form tumors containing neural elements and glandular structures that resemble primitive gut. In culture, these EC cells can be induced to differentiate by exposure to retinoic acid and hexamethylenebisacetamide (HMBA). Differentiation is marked by the disappearance of several cell surface antigens characteristic of human EC cells, and the appearance of other antigens on the various subsets of differentiated derivatives. In retinoic acid-induced cultures, these differentiated derivatives include neurons and cells permissive for the replication of cytomegalovirus, a virus that can cause birth defects in humans. On the other hand, HMBA appears to activate an alternative pathway of differentiation for TERA-2 EC cells, although the identity of the resulting cells remains to be elucidated. In addition to providing a tool for analyzing the evolution of teratocarcinomas in human patients, the TERA-2 EC cells may provide us with insights into the mechanisms of cellular differentiation in the human embryo and a model in which to investigate how teratogenic agents such as HCMV can disrupt these processes.     

Human kinesiology

This paper discusses kinesiological aspects of human locomotion in terms of the incomplete morphological adaptation of the body to current lifestyles. Modern instrumentation used for biomechanical analyses is outlined for the most part in the context of human running. Investigations of spatiotemporal and segmental measures by electronic walkways, photography, optoelectronics and angle-angle diagrams are referred to, as is electromyography and direct force measurement. Three commonly identifiable areas of applied kinesiological research, namely sport, industry and medicine, are considered with reference to the prevention, treatment and after-care of injuries whether received on the sporting field, in the work place, or in an automobile accident.     

Human microcephaly

Microcephaly is defined as a reduction in head circumference and this clinical finding infers that an individual has a significant diminution in brain volume. Microcephaly can be usefully divided into primary microcephaly, in which the brain fails to grow to the correct size during pregnancy, and secondary microcephaly, in which the brain is the expected size at birth but subsequently fails to grow normally. Current work suggests that primary microcephaly is caused by a decrease in the number of neurones generated during neurogenesis, but that in secondary microcephaly it is the number of dendritic processes and synaptic connections that is reduced. Important insights into human neurogenesis are being revealed by the study of rare genetic diseases that involve primary microcephaly, illustrated by the identification of the Microcephalin, abnormal spindle in microcephaly and ataxia-telangiectasia and Rad3-related genes. Furthermore, these findings facilitate the search for the evolutionary changes that have lead to the human brain being so much larger than that of any other primates.     

Human evolution

The common ancestor of modern humans and the great apes is estimated to have lived between 5 and 8 Myrs ago, but the earliest evidence in the human, or hominid, fossil record is Ardipithecus ramidus, from a 4.5 Myr Ethiopian site. This genus was succeeded by Australopithecus, within which four species are presently recognised. All combine a relatively primitive postcranial skeleton, a dentition with expanded chewing teeth and a small brain. The most primitive species in our own genus, Homo habilis and Homo rudolfensis, are little advanced over the australopithecines and with hindsight their inclusion in Homo may not be appropriate. The first species to share a substantial number of features with later Homo is Homo ergaster, or ‘early African Homo erectus’, which appears in the fossil record around 2.0 Myr. Outside Africa, fossil hominids appear as Homo erectus-like hominids, in mainland Asia and in Indonesia close to 2 Myr ago; the earliest good evidence of ‘archaic Homo’ in Europe is dated at between 600–700 Kyr before the present. Anatomically modern human, or Homo sapiens, fossils are seen first in the fossil record in Africa around 150 Kyr ago. Taken together with molecular evidence on the extent of DNA variation, this suggests that the transition from ‘archiac’ to ‘modern’ Homo may have taken place in Africa.