In biology, constructive development refers to the hypothesis that organisms shape their own developmental trajectory by constantly responding to, and causing, changes in both their internal state and their external environment. [1] [2] [3] [4] [5] [6] [7] Constructive development can be contrasted with programmed development, the hypothesis that organisms develop according to a genetic program or blueprint. [3] [8] The constructivist perspective is found in philosophy, most notably developmental systems theory, and in the biological and social sciences, including developmental psychobiology and key themes of the extended evolutionary synthesis. Constructive development may be important to evolution because it enables organisms to produce functional phenotypes in response to genetic or environmental perturbation, and thereby contributes to adaptation and diversification. [6] [8]
At any point in time, an organism's development depends on both the current state of the organism and the state of the environment. The developmental system, including the genome and its epigenetic regulation, responds flexibly to internal and external inputs. One example is condition-dependent gene expression, but regulatory systems also rely on physical properties of cells and tissues and exploratory behavior among microtubular, neural, muscular and vascular systems. [6] [9]
Organisms inherit (i.e., receive from their predecessors) a diverse set of developmental resources, including DNA, epigenetic marks, organelles, enzymes, hormones, antibodies, transcription factors, symbionts, socially transmitted knowledge and environmental conditions modified by parents. [10]
In the course of development, organisms help shape their internal and external environment, and in this way, influence their own development. Organisms also construct developmental environments for their offspring through various forms of extra-genetic inheritance. [11]
No single source of influence has central control over an organism's development. [4] Whilst the genetic influence on development is fundamental, causation does not only occur from the bottom up, but also flows ‘downwards’ [12] from more complex levels of organismal organization (e.g., tissue-specific regulation of gene expression). The result is that many features of organisms are emergent properties that are not encoded in the genome.
Constructive development is manifest in context-dependent gene expression, physical properties of cells and tissues, exploratory behavior of physiological systems and learning.
Although all the cells of an organism contain the same DNA, there can be hundreds of different types of cells in a single organism. These diverse cell shapes, behaviors and functions are created and maintained by tissue-specific gene expression patterns and these can be modified by internal and external environmental conditions.
Assembly of organs, tissues, cells and subcellular components are in part determined by their physical properties. [13] For example, the cell membrane that forms a barrier between the inside and outside of the cell is a lipid bilayer that forms as result of the thermodynamic properties of the phospholipids it's made of (hydrophilic head and hydrophobic tails).
Exploratory processes are selective processes that operate within individual organisms during their lifetimes. [6] [9] In many animals, the vascular, immune and nervous systems develop by producing a variety of forms, and the most functional solutions are selected for and retained, while others are lost. For example, the ‘shape’ of the circulatory system is constructed according to the oxygen and nutrient needs of tissues, rather than being genetically predetermined. Likewise, the nervous system develops through axonal exploration. Initially muscle fibers are connected to multiple neurons but synaptic competition selects certain connections over others to define the mature pattern of muscle innervation. The shape of a cell is determined by the structure of its cytoskeleton. A major element of the cytoskeleton are microtubules, which can grow in random directions from their origin. Microtubule-associated proteins can aid or inhibit microtubule growth, guide microtubules to specific cellular locations and mediate interactions with other proteins. Therefore, microtubules can be stabilized in new configurations that give rise to new cell shapes (and potentially new behaviors or functions) without changes to the microtubule system itself.
In animals, many behaviors are acquired through learning. Social learning and cultural transmission are important sources of adaptive phenotypic plasticity, enabling animals to adapt to their environments even if those environments have not frequently been encountered in the evolutionary history of the species. Social learning also enables stable inheritance of many characters. Cross-fostering of great tit and blue tit chicks show that social learning can result in stable inheritance of species-typical foraging behaviors (foraging height, prey type, prey size, foraging method) as well as nest site choice, alarm calls, songs, and mate choice. [14] [15] Recent killer whale research has demonstrated the divergence of orcas into several species mediated by learned and socially transmitted differences in diets. [16]
Within evolutionary biology, development has been traditionally viewed as under the direction of a genetic program (e.g. [17] ), and metaphors such as genetic ‘blueprint’, ‘program’ or ‘instructions' are still widespread in biology textbooks. [18] By contrast, the constructive development perspective views the genome as a sub-system of the cell shaped by evolution to detect and respond to the signals it receives. [19] These different perspectives affect evolutionary interpretations. One example is the evolutionary significance of environmentally induced phenotypes. Mary Jane West-Eberhard famously suggested that responses to the environment can be the starting point for evolutionary change, [20] termed ‘plasticity-led evolution’. From a programmed development perspective, developmental plasticity is considered to be controlled by genetically specified switches or reaction norms. For these to produce functional responses to environmental change, their reaction norms must have been pre-screened by prior selection. Therefore, ‘plasticity-led evolution’ reduces to the standard evolutionary explanation of natural selection acting on genetic variation. Conversely, ‘plasticity-led evolution’ gains greater significance if development is constructive and open-ended. New functional phenotypes can emerge with little or no initial genetic modification (see facilitated variation [6] [9] ), and provide the new raw material on which natural selection can act (e.g. [21] ).
Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic information of their parents. Through heredity, variations between individuals can accumulate and cause species to evolve by natural selection. The study of heredity in biology is genetics.
Neural Darwinism is a biological, and more specifically Darwinian and selectionist, approach to understanding global brain function, originally proposed by American biologist, researcher and Nobel-Prize recipient Gerald Maurice Edelman. Edelman's 1987 book Neural Darwinism introduced the public to the theory of neuronal group selection (TNGS) – which is the core theory underlying Edelman's explanation of global brain function.
In genetics, the phenotype is the set of observable characteristics or traits of an organism. The term covers the organism's morphology, its developmental processes, its biochemical and physiological properties, its behavior, and the products of behavior. An organism's phenotype results from two basic factors: the expression of an organism's genetic code and the influence of environmental factors. Both factors may interact, further affecting the phenotype. When two or more clearly different phenotypes exist in the same population of a species, the species is called polymorphic. A well-documented example of polymorphism is Labrador Retriever coloring; while the coat color depends on many genes, it is clearly seen in the environment as yellow, black, and brown. Richard Dawkins in 1978 and then again in his 1982 book The Extended Phenotype suggested that one can regard bird nests and other built structures such as caddisfly larva cases and beaver dams as "extended phenotypes".
The genotype–phenotype distinction is drawn in genetics. The "Genotype" is an organism's full hereditary information. The "Phenotype" is an organism's actual observed properties, such as morphology, development, or behavior. This distinction is fundamental in the study of inheritance of traits and their evolution.
Evolutionary developmental biology is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.
A maternal effect is a situation where the phenotype of an organism is determined not only by the environment it experiences and its genotype, but also by the environment and genotype of its mother. In genetics, maternal effects occur when an organism shows the phenotype expected from the genotype of the mother, irrespective of its own genotype, often due to the mother supplying messenger RNA or proteins to the egg. Maternal effects can also be caused by the maternal environment independent of genotype, sometimes controlling the size, sex, or behaviour of the offspring. These adaptive maternal effects lead to phenotypes of offspring that increase their fitness. Further, it introduces the concept of phenotypic plasticity, an important evolutionary concept. It has been proposed that maternal effects are important for the evolution of adaptive responses to environmental heterogeneity.
Niche construction is the process by which an organism alters its own local environment. These alterations can be a physical change to the organism’s environment or encompass when an organism actively moves from one habitat to another to experience a different environment. Examples of niche construction include the building of nests and burrows by animals, and the creation of shade, influencing of wind speed, and alternation of nutrient cycling by plants. Although these alterations are often beneficial to the constructor, they are not always.
In evolutionary biology, the Baldwin effect describes an effect of learned behaviour on evolution. James Mark Baldwin and others suggested that an organism's ability to learn new behaviours will affect its reproductive success and will therefore have an effect on the genetic makeup of its species through natural selection. It posits that subsequent selection might reinforce the originally learned behaviors, if adaptive, into more in-born, instinctive ones. Though this process appears similar to Lamarckism, that view proposes that living things inherited their parents' acquired characteristics. The Baldwin effect only posits that learning ability, which is genetically based, is another variable in / contributor to environmental adaptation. First proposed during the Eclipse of Darwinism in the late 19th century, this effect has been independently proposed several times, and today it is generally recognized as part of the modern synthesis.
The theory of facilitated variation demonstrates how seemingly complex biological systems can arise through a limited number of regulatory genetic changes, through the differential re-use of pre-existing developmental components. The theory was presented in 2005 by Marc W. Kirschner and John C. Gerhart.
Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompasses all types of environmentally induced changes that may or may not be permanent throughout an individual's lifespan.
Origination of Organismal Form: Beyond the Gene in Developmental and Evolutionary Biology is an anthology published in 2003 edited by Gerd B. Müller and Stuart A. Newman. The book is the outcome of the 4th Altenberg Workshop in Theoretical Biology on "Origins of Organismal Form: Beyond the Gene Paradigm", hosted in 1999 at the Konrad Lorenz Institute for Evolution and Cognition Research. It has been cited over 200 times and has a major influence on extended evolutionary synthesis research.
Canalisation is a measure of the ability of a population to produce the same phenotype regardless of variability of its environment or genotype. It is a form of evolutionary robustness. The term was coined in 1942 by C. H. Waddington to capture the fact that "developmental reactions, as they occur in organisms submitted to natural selection...are adjusted so as to bring about one definite end-result regardless of minor variations in conditions during the course of the reaction". He used this word rather than robustness to consider that biological systems are not robust in quite the same way as, for example, engineered systems.
Evolutionary developmental psychology (EDP) is a research paradigm that applies the basic principles of evolution by natural selection, to understand the development of human behavior and cognition. It involves the study of both the genetic and environmental mechanisms that underlie the development of social and cognitive competencies, as well as the epigenetic processes that adapt these competencies to local conditions.
Genetic assimilation is a process described by Conrad H. Waddington by which a phenotype originally produced in response to an environmental condition, such as exposure to a teratogen, later becomes genetically encoded via artificial selection or natural selection. Despite superficial appearances, this does not require the (Lamarckian) inheritance of acquired characters, although epigenetic inheritance could potentially influence the result. Waddington stated that genetic assimilation overcomes the barrier to selection imposed by what he called canalization of developmental pathways; he supposed that the organism's genetics evolved to ensure that development proceeded in a certain way regardless of normal environmental variations.
Developmental systems theory (DST) is an overarching theoretical perspective on biological development, heredity, and evolution. It emphasizes the shared contributions of genes, environment, and epigenetic factors on developmental processes. DST, unlike conventional scientific theories, is not directly used to help make predictions for testing experimental results; instead, it is seen as a collection of philosophical, psychological, and scientific models of development and evolution. As a whole, these models argue the inadequacy of the modern evolutionary synthesis on the roles of genes and natural selection as the principal explanation of living structures. Developmental systems theory embraces a large range of positions that expand biological explanations of organismal development and hold modern evolutionary theory as a misconception of the nature of living processes.
Gerd B. Müller is an Austrian biologist who is emeritus professor at the University of Vienna where he was the head of the Department of Theoretical Biology in the Center for Organismal Systems Biology. His research interests focus on vertebrate limb development, evolutionary novelties, evo-devo theory, and the Extended Evolutionary Synthesis. He is also concerned with the development of 3D based imaging tools in developmental biology.
Transgenerational epigenetic inheritance is the transmission of epigenetic markers and modifications from one generation to multiple subsequent generations without altering the primary structure of DNA. Thus, the regulation of genes via epigenetic mechanisms can be heritable; the amount of transcripts and proteins produced can be altered by inherited epigenetic changes. In order for epigenetic marks to be heritable, however, they must occur in the gametes in animals, but since plants lack a definitive germline and can propagate, epigenetic marks in any tissue can be heritable.
Epigenetics is the study of changes in gene expression that occur via mechanisms such as DNA methylation, histone acetylation, and microRNA modification. When these epigenetic changes are heritable, they can influence evolution. Current research indicates that epigenetics has influenced evolution in a number of organisms, including plants and animals.
The Extended Evolutionary Synthesis (EES) consists of a set of theoretical concepts argued to be more comprehensive than the earlier modern synthesis of evolutionary biology that took place between 1918 and 1942. The extended evolutionary synthesis was called for in the 1950s by C. H. Waddington, argued for on the basis of punctuated equilibrium by Stephen Jay Gould and Niles Eldredge in the 1980s, and was reconceptualized in 2007 by Massimo Pigliucci and Gerd B. Müller.
In biology, reciprocal causation arises when developing organisms are both products of evolution as well as causes of evolution. Formally, reciprocal causation exists when process A is a cause of process B and, subsequently, process B is a cause of process A, with this feedback potentially repeated. Some researchers, particularly advocates of the extended evolutionary synthesis, promote the view that causation in biological systems is inherently reciprocal.