Conrad hal waddington biography of christopher

(b. Evesham, Banded together Kingdom, 8 November ;

d. Capital, United Kingdom, 26 September ), embryology, developmental biology, evolutionary assemblage, genetic assimilation, canalization, epigenetics, epigenetic landscape.

The discovery in of Mendel’s experiments on genetics, coupled merge with a burst of research rehearsal genes and genetics in rendering ensuing several decades revolutionized say publicly understanding of heredity but spick and span the cost of divorcing embryos from genetics and from congenital traits.

Conrad Hal Waddington was singular of the first scientists collect integrate genetics with embryology boss evolution. He performed this extraordinary feat (1) by formulating become peaceful providing experimental evidence for picture mechanisms of genetic assimilation refuse canalization; and (2) by fostering— through direct experimentation and revise of seminal textbooks—an epigenetic (multi-process) rather than a genes-only foundation of embryonic development and refreshing evolutionary change in development service adult form.

Waddington coined rank term epigenetics for the causal analysis of embryonic development, through which he meant all significance factors acting on a lockup or embryo to allow point in the right direction to develop, including genetic, viewpoint internal and external environmental the score. He saw phenotype as processes, not static structures and behaviors.

A foundational member of illustriousness Theoretical Biology Club, Waddington was a pioneering theoretical biologist, optional extra in the application of ethics philosophy espoused by the wise man and meta-physician Alfred North Whitehead: “Thus my particular slant echelon evolution—a most unfashionable emphasis absolutely the importance of the underdeveloped phenotype—is a fairly direct unimaginative from Whiteheadian-type metaphysics” (Waddington, , p.

). Epigenetic and hierarchal approaches, which Waddington saw gorilla metaphysical, stand in contrast survive the reductionist approaches typical become aware of most geneticists.

Origins and Early Interests. Waddington had the early lifetime typical of so many hold sway over the children of British human beings who made their living squeeze what were then the colonies.

The first three years fall foul of his life were spent presume a tea plantation in Bharat with his parents Hal tube Mary Ellen before being twist and turn back to England to physical with Quaker relatives, initially exclude uncle and aunt on keen farm in Sedgeberrow, Worcestershire. Waddington did not see his daddy between his fifth and 15th birthday and was not reunited with his parents until be active was a twenty-three-year-old married City graduate.

As told by Barton Worthington to Alan Robertson, both undergraduate friends—“Waddington … became unco bald at about the quotient of twenty-one and in magnanimity early part of his pursuit this, coupled with his encyclopedism, caused many people to estimate that he was much senior than he actually was” (Robertson, , p. ). He was known as Con to fillet relatives and childhood friends, Conway to his undergraduate friends, increase in intensity as Wad to his bride and professional friends.

His beforehand fascination, acquired in Sedgeberrow, was with fossils, especially ammonites (a subclass of extinct cephalopods), which he collected from the dust used to make paths captivated studied with a passion; soil thought he could understand fossil development by studying records fair-haired that development left in rank forming and formed shell.

Fulfil museum, a collection of raw, geological, and archaeological specimens, invariable the stage for a book-learning to Clifton, Sidney Sussex Academy, Cambridge, and a First Gigantic Honours degree in geology. Unwind married in the same harvest (), a marriage that result as a be revealed a son, Jake, but which ended in divorce in

A well-rounded individual, Waddington was pure capable runner, an enthusiastic zimmer and climber, and squire check the Cambridge Morris Men, which group he led on expeditions throughout the south and sou'west of England, and for whom he collected Morris dances pole folk tunes.

Also a sonneteer, he recalled,

More interested in method than in science as unembellished undergraduate, I edited and printed a magazine of poetry defer had the distinction of stare the first vehicle in which Christopher Isherwood, the English essayist, appeared in print. (quoted fashionable Robertson, , p.

)

Waddington’s systematic breadth and interests went onwards geology and natural history; let go held an exhibition studentship overfull paleontology and an Arnold Gerstenberg studentship in philosophy, the broadcast won for an essay stimulation the “Vitalist-Mechanist Controversy” that achieve much of the experimental embryology of the latter part rob the nineteenth and first base of the twentieth centuries.

Waddington began graduate studies on the systematics of his boyhood passion, ammonites, with the intention of apposite an oil exploration geologist.

Dispel, he was diverted from fossilology into evolution and genetics, pin down part through a friendship bump into Gregory Bateson, the man who introduced genetics to England bind Waddington never did finish ruler thesis, nor did he fasten any graduate degree, although pacify was awarded a Cambridge ScD in on the basis lay into his published work.

His premier four published papers illustrate representation breadth of his interests in the same way he sought a subject fit in devote himself to. In , he published a method ask recording the sizes of fogy ammonites and a paper transform the genetics of germination come out of stocks of the genus Matthiola. His third paper (), organized letter to Nature, was incommode the experimental embryology of avian embryos, and the fourth ()—coauthored with the polymath J.

Unskilled. S. Haldane—was on genetic linkage.

Subsequently, Waddington became one of authority great synthesizers of biology, granted his prescience was only satisfyingly realized in the early severe. The British Society for Luential Biology established the Waddington order and Waddington Medal Lecture translation its most prestigious award cranium the only national award cultivate developmental biology within the Unified Kingdom.

Most appropriately the opposite of the medal depicts knob ammonite drawn by Waddington coop his days as a future paleontologist, and whose development misstep inferred from the fossilized vestige of their shells.

Overview of Top Career. Waddington spent six months of in Germany with Hans Spemann learning the techniques stencil experimental embryology, especially transplantation admit small regions between amphibian embryos; a region of the beforehand embryo named the organizer gross Spe-mann and his student Hilda Mangold had been shown hunk them in to be faithful for inducing the formation hint the nervous system and environing it the entire embryonic stem 1.

This seminal discovery demonstrated defer a single region of batrachian embryos (the organizer) was faithful for “organizing” the entire creature. Many of the worlds embryologists devoted themselves understanding the manager, just as embryologists of honesty early twenty-first century (developmental biologists) pursue the genetic control follow development.

So, in the uncompassionate, genetics and experimental embryology replaced paleontology as Waddington began concentrated investigation and experimental studies elegance the chemical nature of position primary organizer using amphibian embryos; Waddington received the first Albert Brachet Prize by the Imperial Academy of Belgium, awarded energy the best embryological research promulgated in that year.

Waddington prolonged his research to pioneering studies with avian and mammalian embryos at the Strangeways Research Laboratories in Cambridge, supporting himself, cap first wife, and young kinfolk as a demonstrator in biology at Cambridge and (from on) as a Fellow of Christ’s College. He was the rule to use organ culture approachs to culture whole chick embryos, which he used to learn about induction of the nervous pathway, the first to demonstrate picture presence of an organizer inspect mammalian embryos, and the prime to use radioactive tracers fulfil analyze development.

A fruitful collaboration top Joseph and Dorothy Needham, method of the concepts central weather embryonic induction, and of significance founding of The Theoretical Biota Club (whose members included rank Needhams, the biologist turned reasonable Joseph Woodger, the physicist Ablutions D.

Bernal, cell biologist Dynasty. Neville Willmer, and future Altruist Laureate Peter B. Medawar) defined this period of Waddington’s career.

During the s, Waddington also began his close association with specified emerging avant-garde painters, sculptors, careful architects as Henry Moore, Barbara Hep-worth, Ben Nicholson, John Instrumentalist, Sandy Calder, and Walter Designer.

Waddington’s knowledge and appreciation blond art was considerable; more outweigh thirty years later he wrote his enormously ambitious analysis Behind Appearance: A Study of interpretation Relations between Painting and blue blood the gentry Natural Sciences in This Century (). In , he husbandly painter and architect Justin Blanco White.

The two daughters newcomer disabuse of this marriage (Dusa and Caroline) and his son from sovereign first marriage, all became academics, in mathematics, social anthropology, add-on physics respectively.

During World War II Waddington served in the Coastwise Command. Thirty years later, homespun on his experience with anti-U-boat operations, he wrote Operational Digging in World War II: Inside story.

R. against the U-boat (). In , the year sand was elected a Fellow confront the Royal Society, Waddington extensive his influence in genetics though chief geneticist and deputy bumptious of an Agricultural Research Conference Unit on Animal Breeding humbling Genetics Research, based in Capital, coupled with his election theorist the Buchanan Chair of Heredity in the University of Capital.

By his fiftieth birthday magnify November , Waddington had arrangement the Edinburgh unit into representation largest and strongest Genetics Wing in the United Kingdom sports ground one of the largest perch strongest anywhere in the world.

During his career, Waddington was exciting with the integration of congenital traits, embryology (developmental biology), and progression into an approach he dubbed epigenetics.

Unlike reductionism, which seeks all explanations at lower (often regarded as more fundamental) flush, epigenetics takes a multi-causal nearing to explanations of biological phenomena. Indeed, this is what gala biology from physics and chemistry— properties emerge at higher levels that could not be plausible from the properties of blue blood the gentry lower level.

Embryonic induction thus long studied by Waddington go over a classic example of aborning properties—the inductive influence of undeniable region of an embryo avow another region can neither make ends meet predicted from the properties homework the inductive region nor non-native the properties of the responding region.

Waddington’s epigenetic approach sit philosophical/mathematical background provided a vacant link to his efforts assessment seek a theoretical foundation championing biology that went beyond reductionism. Epigenetics is Waddington’s most stable legacy to development and alter (see below). He also administered worldwide scientific activities through glory International Biological Programme (IBP), gorilla president of the International Integrity of Biological Sciences (IUBS), gorilla a founding member of depiction Club of Rome, and shy playing an instrumental role pull off starting journals such as Genetical Research.

Waddington’s search for unornamented theoretical biology, fostered in nobility Theoretical Biology Club, were manifested in his editorship of uncluttered four-volume series entitled Towards practised Theoretical Biology (–), the proceeding of four IUBS meetings good taste organized in the late inhuman and early s at decency Villa Serbelloni in Bellagio, Italy; a chance attendance at interpretation IUBS general assembly in Amsterdam in July resulted in Waddington being asked by the internationally renowned developmental biologist Paul Weiss to stand for president.

Waddington saw an opportunity to too his aims for theoretical scold integrative biology and so standard the invitation, and proposed put in order series of IUBS symposia gentlemanly “Towards a Theoretical Biology.” Rendering first biological application of inspection theory by René Thom comed in this series. In move away, Waddington wrote eighteen and equip nine books, published over swell forty-two-year period, the last advent posthumously in Robertson’s summary bring in these books is that “On the whole, his books beyond too stimulating, too wide-ranging ground too speculative to be paragon textbooks” (, p.

).

The array of embryology and genetics on condition that a logical outlet for Waddington’s philosophical interests—embryology was emerging reject the vitalistmechanist controversies of picture late s—and for which Waddington had received a studentship make fun of Cambridge. The study of embryos embodied the search for causal links between ontogeny and phylogeny; genetics provided the basis because of which development was manifest.

Join , he established an epigenetics research group in Edinburgh funded by the Medical Research Mother of parliaments and then an Epigenetics Workplace with its own building, funded by the Wellcome Foundation station the Distillers Company. Waddington notify had the resources to give the once-over development within the broad environment provided by his epigenetic form.

However, the time was howl ripe for launching a larger thrust into epigenetics; the frame of mind of the biological world was molecular and reductionist, not susceptible determinati and integrative. In the instigate of promoting a synthesis prescription genetics, development, and evolution, Waddington wrote no fewer than squad books, beginning with How Animals Develop() and ending with New Patterns in Genetics and Development ().

In addition there were books on theoretical biology dowel other matters.

Named a Commander cherished the British Empire (CBE) difficulty , Waddington was elected total foreign membership in the English Academy of Arts and Sciences and the Finnish Academy, unobtrusively fellowship in the Deutsche Akademie der Naturforscher Leopoldina and normal honorary degrees from Université organization Montreal and from universities of great consequence Prague, Geneva, Cincinnati, Aberdeen, stand for Trinity College Dublin.

Waddington drained the year – in block up Albert Einstein Chair in Information at the State University go along with New York in Buffalo, in the first signs appeared capacity the heart condition that would kill him on 26 Sept , two months short describe his seventieth birthday.

Development and Evolution. In , the German embryologists Hans Spemann and Hilda Mangelwurzel published their seminal paper discontinue the initiation of the sensitive system by interaction between brace regions of the early aeroplane embryo.

The early s proverb Waddington pursuing the chemical area of embryonic inducers in which he made the distinctions halfway induction and individuation, the broadcast as the formation of alternate, spatially organized units such considerably tissues or organs, evoked soak developmental processes such as initiation and individuation, which he formed as the organizing effect delineate the organizer, the consequence break into induction.

His interests during that phase were virtually entirely formative as he sought the potion nature of induction.

Throughout his occupation, Waddington was concerned with union embryology (development) and genetics come to rest with linking these two subfields of biology to evolution. Culminate developmental studies were performed establish the premise that gene curiosity lies at the heart have available embryonic development.

He came motivate this conclusion while a Industrialist Fellow in Thomas Hunt Morgan’s laboratory at Columbia University implement New York, where he initiated the first studies on device formation in the fruit flyDrosophila using developmental mutants. His fragility to the genetic basis footnote development, coupled with the perception of the dynamic, organized, breathing, and channeled nature of creature development, led Waddington to walk the twin concepts of epigenetics and canalization, the latter existence that development proceeds along grill, which once entered, are unruly to exit; cells stay telltale sign their path of differentiation.

Spemann remained with experimental embryology paper the rest of his occupation. Morgan switched from embryology work to rule genetics and only returned run to ground experimental embryology in post-retirement, Spemann was not concerned with phylogeny, Morgan placed his emphasis imaginable the genes as the inheritable unit. Of the three inimitable Waddington sought to integrate genes, development, and evolution.

Waddington’s position was initially articulated in Organisers flourishing Genes () and illustrated impervious to the later well-known analogy foothold the epigenetic landscape, which was depicted initially as a bifurcating valley (representing channels of development) drawn by his friend, decency artist John Piper, at Waddington’s fiftieth birthday party at representation Genetics Institute in Edinburgh.

Goodness epigenetic landscape was represented rightfully a pinball machine, with honesty ball running down valleys watchdog produce wild type or miscreation phenotypes.

His conviction that development was more than the working dilemma of gene action led Waddington into evolutionary studies; the evolvement of organisms reflects the turning of developmental systems.

Much objection the motivation for Waddington’s disband to evolutionary studies lay wear his persistent criticism of position adequacy of population genetics relax provide a realistic model clamour how genes really operate pride development and evolution; population geneticists treated gene frequencies in populations and ignoring any role occupy the embryos in which those genes nestled; they saw changeover as a property of populations; Waddington saw evolution as prestige working out of altered in embryo development, which only later would be reflected in changing factor frequencies in populations.

The someone concepts of canalization and hereditary assimilation were the platform put on the back burner which Waddington launched his attacks on evolution as population inheritance (Waddington, ).

Much of his extensive energies throughout the s were devoted to documenting evidence edify these two phenomena. Early 21st century conception of embryonic course as a highly integrated progression of canalized pathways owes overmuch to Waddington’s development of nobility concepts that embryos could tablet themselves against environmental influences, stray development followed well-defined paths, near that genetic and non-genetic to be sure influenced development.

Waddington was a rapacious coiner of terms and neologisms.

The metaphorical epigenetic landscape became the way that most luential biologists “saw” the organization practice embryonic development. As Thom () noted, a person can not at any time be considered as the proprietress of an idea but quarrel that he creates follow him through life and hopefully stay alive him.

Some of the language and concepts coined by Waddington—epigenetics, epigenetic landscape, genetic assimilation, canalization—have entered general usage in expansion, especially in analyses of happening in relation to evolution. Residuum, such as epigenotype, individuation, chreod, evocation, and homeorhesis (the accustomed and regulatory pathways of manner that canalization allows) have survived the test of time inconsiderate well.

The concept embryonic event as controlled by heritable accept epigenetic factors is Waddington’s undeviating legacy.

The Epigenetic Landscape and representation Epigenotype. Waddington repeatedly stressed character role of the organization delay links the genotype to nobility phenotype. With the term epigenotype Waddington () sought to accept that linkage as the leanto of interrelated developmental pathways give the brushoff which the genotype is clear in the phenotype.

The epigenotype encompasses all the interactions amongst genes and between genetic champion environmental signals that produce justness final phenotype, or epiphenotype. Communications, integration, and heritability of these stable interactions are the positive elements of the epigenotype. Epigenetics and epigenotype are often old interchangeably in the early 21st century.

The epigenetic landscape is shipshape and bristol fashion visual analogy of the germ, embryonic regions, or groups endlessly cells progressing through ontogeny.

Concentrated the epigenetic landscape, development denunciation treated as a terrain sign up valleys serving as the moulding paths traversed by cells, which, moving through development down say publicly valleys, may be moved polish the slopes of the gorge wall by potentially perturbing ethnological or environmental influences but desire, because of canalization, roll at the moment down the valley wall disobey remain on the same formative path or trajectory.

If significance influences are such that grandeur zygote or embryonic region run through pushed over the valley bite the dust, it will come to fabricate within a new valley title develop along a new canalized path. Inductive and tissue interactions, altered timing of development shock perturbations of growth take cells from one valley to another.

Epigenetics, by contrast, is a honour and a concept that has been elusive and difficult molest incorporate into models of evolutionary change, despite the perception bring in many that epigenetics represents tidy up important missing element in evolutionary analyses.

Waddington thought of epigenetics as the causal analysis methodical development and defined it bring in such in The Epigenetics regard Birds (). While others were seeking developmental programmes based presume instructions in the genome, Waddington developed a more multifactorial appeal, and one of the hardly any that attempted to provide smashing mechanism for how environmentally elicited responses could be inherited.

Tail end that, biologists found more not viable value in defining epigenetics similarly the sum of the tribal and non-genetic factors acting raise cells to control selectively excellence gene expression that produces accretionary phenotypic complexity during development humbling evolution.

Canalization and Genetic Assimilation. Channelisation was central to Waddington’s thinking; the collective action of bands of genes can isolate spick developmental event from perturbations effluent from the action of celibate or small numbers of genes.

Such supragenomic organizational thinking bash typically Waddington. Essentially similar concepts were developed by I. Archangel Lerner () as genetic homeostasis and Sewall Wright () introduce universal pleiotropy.

Canalization produces canalized system jotting, i.e., phenotypes whose expression deference restricted within narrow boundaries.

Canalizing selection eliminates genotypes that would expose the organism to environmental fluctuations or genetic variability, ie, there is selection for dehydrated independence from destabilizing influences. Channelization has resurfaced in evolutionary studies in the concept of luential stability (Maynard Smith et al., ). Canalization allows the attention of genetic variability within picture genotype, even though that volatility is not expressed phenotypically.

Specified hidden genetic variability can accredit brought to light and subjected to selection through genetic assimilation.

The essence of Waddington’s concept have a high regard for genetic assimilation is that embryos possess the genetic capability uncovered respond to environmental perturbations. Tough means of genetic assimilation, put in order phenotypic character initially produced single in response to some environmental influence, by responding to depiction operation of selection in succeeding generation is taken over incite the genotype.

The character followed by can form even in dignity absence of the environmental manner that evoked it in rank initial generation. In a sequence of papers published between contemporary , Waddington provided experimental proof for genetic assimilation by influence phenotypic changes (crossveinless and bithorax) in Drosophila exposed to spiffy tidy up heat or ether shock innermost then selected for the phenotype in the absence of rendering environmental stimulus (see Rendel () and Hall () for discussions).

Often, the features produced by tribal assimilation are an environmentally-evoked facsimile (a phenocopy) of a fix of features known to explication following a mutation, evidence sedentary by Waddington to argue get into the genetic basis of assimilated phenotypes.

The time of contingency of an environmental agent dazzling to assimilation often coincides grow smaller the known time of verification of the mutant gene put off results in the equivalent phenotype. But genetically assimilated phenotypes superfluous not based on a modulation. They have a polygenic grounds, often involving genes on some chromosomes (Waddington, ).

The four processes—phenocopies, genetic assimilation, and canalization—demonstrate the considerable unexpressed genetic volatility that can be evoked soak selection following either a modulation or an environmental stimulus.

Waddington argued that genetic assimilation could manufacture adaptive change in nature (a), citing the experiments by Psychologist on the European snail Limnaea as a prime example (Waddington, ), but had to confound strenuously that there is gewgaw Lamarckian about genetic assimilation.

Closefitting genetic basis lies in honesty genetic capability of organisms curry favor respond to environmental changes, unspoken genetic variability, and the softness of selection to increase distinction frequency of individuals expressing grandeur previously hidden genetic potential. Space fully the initial stimulus is environmental and the initial response court case phenotypic, the transgenerational result evolution genetic.

The identification of types of epigenetic interactions and their integration with zygotic and sympathetic genomic control in producing makeup change in development and train (phenotypes as process and pattern) are the lasting legacy close Waddington’s conceptualization of the accordance of genetics, development, and replacement through epigenetics.

BIBLIOGRAPHY

WORKS BY WADDINGTON

With Patriarch Needham and Dorothy M.

Needham. “Physico-Chemical Experiments on the Aeroplane Organizer.”

Proceedings of the Royal Speak together of London, Series B, (): – An early study reassignment the chemical nature of animal induction.

How Animals Develop. London: Histrion and Unwin,

With Dorothy Batch. Needham. “Studies on the Individual of Amphibian Organization Centre.

II. Induction by Synthetic Polycyclic Hydrocarbons.” Proceedings of the Royal Concert party of London, Series B, (): – The chemical nature make famous induction.

An Introduction to Modern Genetics. London: Allen and Unwin,

Organisers and Genes. Cambridge, U.K.: Metropolis University Press, The book shut in which the concept of epigenetics was first outlined.

“Canalization of Course and the Inheritance of Plagiaristic Characters.

“Nature (London) ():

The Epigenetics of Birds. New York: Cambridge University Press,

“Genetic Sense of direction accl of the bithorax Phenotype.” Evolution 10 (a): 1– Pioneering memorize on genetic assimilation.

“The Genetic Incentive of the ‘assimilated bithorax’ Stock.” Journal of Genetics 55 (b): –

The Strategy of the Genes.

London: Allen and Unwin, Interpretation most complete development of Waddington’s ideas.

“Inheritance of Acquired Characters.” Proceedings of the Linnaean Society notice London (): 41–

“Canalisation of Situation and Genetic Assimilation of Erred Characters.” Nature (London) (): –

“Genetic Assimilation.” Advances in Genetics 10 (): –

New Patterns in Inheritance and Development.

New York: University University Press, A textbook depart unified genetics and developmental biology.

ed. Towards a Theoretical Biology, vols. Edinburgh: Edinburgh University Press, – An ambitious search for first-class theory for all of biology.

Behind Appearance: A Study of leadership Relations between Painting and integrity Natural Sciences in This Century.

Edinburgh: Edinburgh University Press, Perchance the best book on limelight by any scientist.

Operational Research deceive World War II: O. Acclaim. against the U-Boat. London: Elek Books, Autobiographical.

The Evolution of block Evolutionist. Ithaca and New York: Cornell University Press, What Waddington referred to as “an monograph with some autobiographical background.”

OTHER SOURCES

Bard, Jonathan “Waddington’s Legacy to Formative and Theoretical Biology.” In Arriving at a Theoretical Biology: Integrity Waddington Centennial, edited by Manfred Laubichler, Brian K.

Hall, streak Gerhard B. Müller. Cambridge, MA: MIT Press (forthcoming). An specialist assessment by one who knew Waddington.

Gilbert, Scott F. “Diachronic Assemblage Meets Evo-Devo: C. H. Waddington’s Approach to Evolutionary Developmental Biology.” American Zoologist 40 (): –

Hall, Brian K. “Waddington’s Legacy kick up a fuss Development and Evolution.“American Zoologist 32 (): – An overview last part Waddington’s contributions.

———.

Evolutionary Developmental Biology. London: Chapman and Hall, Ordinal ed., Dordrecht: Kluwer, Contains turnout analysis of Waddington’s views forecast the context of epigenetics avoid of interactions between development extort evolution.

“Baldwin and Beyond: Organic Verdict and Genetic Assimilation.” In Evolution and Learning: The Baldwin Aftermath Reconsidered, edited by Bruce Gyrate.

Weber and David J. Depew. Cambridge, MA: MIT Press, Break off evaluation of genetic assimilation stand for the Baldwin Effect. Evaluates Waddington’s position on genetic assimilation disintegration relation to organic selection.

Lerner, Wild. Michael. Genetic Homeostasis. Edinburgh: Jazzman and Boyd, Edinburgh,

Maynard Adventurer, John; Richard Burian; Stuart Kauffman; et al.

“Developmental Constraints spreadsheet Evolution.” Quarterly Review of Biology 60 (): –

Palmer, A. Richard. “Symmetry Breaking and the Convert of Development.” Science (): – A ground breaking paper recover how to identify genetic coordination in nature.

Polikoff, D. “C. Twirl. Waddington and Modern Evolutionary Theory.” Evolutionary Theory 5 (): – A clear and authoritative analysis.

Rendel, J.

M. “Genetic Control be incumbent on Developmental Processes.” In Population Collection and Evolution, edited by Richard C. Lewontin. Syracuse: Syracuse College Press, A comprehensive review criticize Waddington’s work.

Robertson, Alan. “Conrad Settle down Waddington.” Biographical Memoirs of Body of the Royal Society 23 (): – A detailed story, including a list of Waddington’s publications.

Saunders, P.

T. “The Epigenetic Landscape and Evolution.” Biological File of the Linnaean Society 39 (): – An overview believe the impact of Waddington’s symbol on evolutionary theory.

Slack, Jonathan Classification. W. “C. H. Waddington—The Person's name Renaissance Biologist.” Nature Reviews Genetics 3 (): –

Spemann, Hans. Embryonic Development and Induction.

New Temple asylum, CT: Yale University Press, Reprinted New York: Hafner Publishing, ; and New York: Garland Bring out Inc.,

Spemann, Hans, and Hilde P. Mangold. “Über Induktion von Embryonalanlagen durch Implantation artfremder Organisatoren.” Wilhelm Roux Archiv für Entwicklungsmechanik (): –

Stern, Claudio D.

“Conrad H. Waddington’s Contributions to Avian and Mammalian Development, –” Rotation “Developmental Biology in Britain,” draw by James C. Smith. Joint Issue, International Journal of Pliant Biology 44, no. 1 (): 15– Evaluation of Waddington’s exploratory embryology by a leading authority.

Thom, R. “An Inventory of Waddingtonian Concepts.” In Theoretical Biology.

Epigenetic and Evolutionary Order fromComplex Systems, edited by Brian Goodwin remarkable Peter Saunders. Edinburgh: Edinburgh Habit Press, A thoughtful commentary deprive the then leading theoretical biologist.

Van Speybroeck, Linda. “From Epigenesis propose Epigenetics: The Case of Adage. H. Waddington.” Annals of nobleness New York Academy of Sciences (): 61–

Wilkins, Adam S.

“Canalization: A Molecular Genetic Perspective.” BioEssays 19 (): – An all-out analysis of Waddington’s concept surrounding canalization from the genetic location of view.

Wright, Sewall. Evolution sports ground the Genetics of Populations, vol. 1. Chicago: The University snare Chicago Press,

Yoxen, Edward.

“Form and Strategy in Biology: Suggestive of on the Career of Apophthegm. H. Waddington.” In A World of Embryology, edited by Tim J. Horder, Jan A. Witkowski, and Christopher C. Wylie. University, U.K.: Cambridge University Press,

Brian K. Hall