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The Ontogeny of Information
Developmental Systems and Evolution
By Susan Oyama Duke University Press
Copyright © 2000 Duke University Press
All rights reserved.
ISBN: 978-0-8223-8066-5
CHAPTER 1
Introduction
In the Western religious tradition, God created the world by bringing order to chaos. By imposing form on inchoate matter, he acted according to a convention that was very old indeed, one that separated form from matter and considered true essence to reside in the former. (Interestingly enough, "form" is cross-referenced to "universals" in the Encyclopedia of Philosophy.) This separation of form from matter underlies and unites all the versions of the nature-nurture antithesis that have so persistently informed our philosophical and scientific approaches to the phenomena of life. That it is not a necessary one is evidenced by studies of creation myths from other cultures (G. Bateson, 1972; Long, 1963).
Those who have argued over the origin of ideas and of biological beings have usually agreed that form in some sense preexists its appearance in minds and bodies. They have only disputed the method and time of its imposition. Most solutions to the puzzle of how form arises, therefore, including the most recent biological dogma, have incorporated the assumption that form is to be explained by pointing to a prior instance of that very form. To the extent that this is true, they are of limited value in answering questions about origins and development. Whether it is God, a vitalistic force, or the gene as Nature's agent that is the source of the design of living things and that initiates and directs the unfolding of the design thus matters little to the structure of the argument. Nor are the problems inherent in such a notion lessened by the use of a succession of metaphors, such as genetic plans, knowledge, and programs, to serve these cognitive and intentional functions.
In an increasingly technological, computerized world, information is a prime commodity, and when it is used in biological theorizing it is granted a kind of atomistic autonomy as it moves from place to place, is gathered, stored, imprinted, and translated. It has a history only insofar as it is accumulated or transferred. Information, the modern source of form, is seen to reside in molecules, cells, tissues, "the environment," often latent but causally potent. It is thought to enable these molecules, cells, and other entities to recognize, select, and instruct each other, to construct each other and themselves, to regulate, control, induce, direct, and determine events of all kinds. When something marvelous happens, whether it be the precise choreography of an "instinctive" behavior or the formation of an embryonic structure, the question is always, Where did the information come from? Was it already in the animal or the developing tissue, or did it have to be put in through learning or perhaps some embryological organizer? Was selection or instruction responsible? Is this a phylogenetic or an ontogenetic adaptation? (Was the information acquired by the species or must it have been acquired through individual experience?) The ease with which extreme nature and nurture positions are parodied ensures that no one will stand behind either straw man. No one really argues, that is, either that livers and ideas are literally in the cell or that organisms are devoid of structure, pristine pages on which anything at all may be written (and even a page has structure!). Or, to put it in negative terms, no one contends that either developmental conditions or the genes are totally irrelevant to development. Any locution that dissociates one from the straw man even minimally, however, seems to offer protection from criticism. "Encoded potential" and "biological constraints," then, allow everyone to return to work, the pesky conceptual issues behind them, peace apparently restored.
But conceptual issues are not detachable from empirical ones. They are there whenever one decides what to count as data, interprets them, draws theoretical or practical inferences from them, and frames the next research question in terms of a particular method. Meanwhile disagreement is not truly resolved, even though a generally interactionist vocabulary is rapidly becoming universal. The failure of this shared vocabulary to resolve old conflicts is significant; I suggest the reason is that ways of thinking about form, constancy, and change have been left largely undisturbed. (See presentations by, and commentary on, Eibl-Eibesfeldt, 1979.) At present, all that needs to be pointed out is that these exchanges often suggest, whatever the other philosophical biases of the speaker, a "preformationist" attitude toward information: it exists before its utilization or expression. Some views allow assembly of information from a variety of sources, but this in turn implies that it exists in several loci before being collected;such views thus perpetuate the mistake while seeming to correct it. In addition, information is conceived to be a special kind of cause among all the factors that may be necessary for a phenomenon, the cause that imparts order and form to matter. The alternative to such a preformationist attitude toward form is not a classical epigenetic one. Not only did this traditionally require that order arise from chaos, an unsatisfactory solution at best, but it often posited a vitalistic force as well, to effect the recurrent miracle. This brings us full circle to the preexisting form, this time ready to inform the formless rather than simply waiting to unfold. Instead, it is ontogenesis, the inherently orderly but contingent coming into being, that expresses what is essential about the emergence of pattern and form without trapping us in infinite cognitive regress (where was the pattern before it got here?). A proper view of ontogeny, however, that doesn't simply resolve into one of the old ones, requires that the idea of ontogenesis apply not only to bodies and minds, but to information, plans, and all the other cognitive-causal entities (more on this phrase in the next chapter) that supposedly regulate their development. Developmental information itself, in other words, has a developmental history. It neither preexists its operations nor arises from random disorder. It is neither necessary, in an ultimate sense, nor a function of pure chance, though contingency and variation are crucial to its formation and its function. Information is a difference that makes a difference (G. Bateson, 1972, p. 315), and what it "does" or what it means is thus dependent on what is already in place and what alternatives are being distinguished.
If information (along with its terminological and conceptual variants) is developmentally contingent in ways that are orderly but not preordained, and if its meaning is dependent on its actual functioning, then many of our ways of thinking about the phenomena of life must be altered. Development can no longer be explained as a combination of translated information from the genes (to make innate features) and information acquired from the environment (to modify, supplement, or complete those features). Nor can phenotypic features be divided into those that are programmed or biological and those that are not, or ranged on a continuum of relative degrees of programming, at least not if "programming" is meant to express anything more than relative predictability in a given context.
The association of biology (or, in many discussions, the genes) with the invariant, the intractable, or the inevitable is a serious error and one that must be undone. The further association of nature with biology compounds it. To say this is most emphatically not to deny relative invariance of some aspects: of the phenotype or developmental conservatism in any particular setting. It is to deny that they are the defining features of some special set of processes or characters that may be dubbed "biological" and distinguished from "flexible" ones by the role of genes in their developmental causation. It is further to deny that the common is always necessary. Nor am I disputing the possibility or utility of genetic analysis or of accounting for phenotypic differences by specific environmental or genetic variation; it is the granting of special ontological status to certain traits on the basis of such findings to which I object, for this leads only too easily to conclusions about nature and fate that are neither justified nor useful.
It is my contention that developmental information does "develop," not by special creation from nothingness, but always from the conditional transformation of prior structure—that is, by ontogenetic processes. Depending on the level of analysis, such transformation can be described as resulting from interactions among entities, such as cells or organisms, or from interactions within an entity, such as an embryo or a family. Since sets of interactants can be at least partially nested, a change in focus is sufficient to shift from one perspective to the next. This idea, that "information" has a developmental history, doesn't arise spontaneously from nothing, but is itself prepared by empirical and conceptual developments on many fronts: various "interactionisms" in psychology, aspects of systems theories and of Piagetian genetic epistemology, discoveries in molecular and developmental biology (though not much of the most general level of interpretation of these discoveries), cognitive psychology, and ethology. It reframes some intractable problems in these fields and offers an alternative way of conceptualizing form and causation, change and variability, normality and necessity. The notion of the ontogenesis of information in what I call developmental systems is implicit in much that is said and done in these sciences today, but because it usually remains implicit, it coexists all too easily with its own contradiction. It is my intention to make it fully explicit, but to do so it is necessary also to render explicit the assumptions supporting the nature-nurture complex. Once the assumptions are clear, it becomes easier to see the problems that attend them.
It is not a preference for the implicit, however, that has delayed full formulation of this concept of information, but rather a multitude of factors, including the conviction that scientific Darwinism requires the distinction between some notion of inherited traits or forms and some notion of acquired traits or forms, no matter how the distinction is made, and that the alternative is a lapse into the nether world of nonscientific Lamarckianism. Waddington remarks that "Lamarck is the only major figure in the history of biology whose name has become, to all intents and purposes, a term of abuse" (1975, p. 38). Though I do not intend to go into the question of what Lamarck really meant and what happened to those ideas in the history of biology, I think it is true that he has come to symbolize, partly by association with Lysenko and therefore with everything that is seen to be inimical to progressive science in the enlightened world, much more than he needs to. This symbolic complex has made it difficult for us to synthesize our notions of inheritance with our notions of development. My intention, then, is not to rehabilitate Lamarck or to present a Lamarckian view; scholars disagree on what such a view would be, and I will refer to him almost not at all. I do hope to show that the opposition of inherited or innate characters to acquired ones (as distinct from heritable or non-heritable variance) is based on a misapprehension of what it takes to be a properly scientific Darwinian. "If we're all Darwinians," Symons (1987) asks, "what's the fuss about?" And if we're all interactionists, again, why the fuss? I suggest the fuss arises (and sometimes doesn't arise when it should) because we haven't figured out what either requires, much less how to be both at the same time. A solution that combines encoded nature with varying doses of contingent nurture is no solution at all.
The question of precisely what is passed on in heredity, and therefore the mechanics and meaning of phylogenesis, is inevitably implicated in this conceptual reorganization. The conventional idea, that is, that organs, developmental sequences, and certain behaviors are transmitted in the genes, or that information or rules for those organs, sequences, and behaviors are so transmitted, continues to undergo progressive reformulation as successive versions are criticized and discarded. But, I submit, these changes are necessarily only cosmetic because they are not allowed to do violence to the underlying idea of inherited form or the associated belief in two kinds of development: one to manifest that form, and another for everything else. Thus we have a procession of nature-nurture oppositions based on this developmental dualism, almost as though we were generating the norm of reaction for this central concept of genetic information.
One of the problems I will address, then, is the idea of developmental information in the chromosomes. What does it explain? What is it information about? What important questions does it or does it not clarify? When is it invoked metaphorically, and what could it mean to treat it literally? When, if ever, does it really help us understand how the living world is constituted and what the prospects for that world are? Intimately related is the problem of the meaning of the "biological." Insofar as "biological bases" have replaced innate knowledge and structure in the vocabularies of many, they have inherited, so to speak, the implications of the earlier terms while legitimating them with the aura of methodological sanctity. But what does it mean for something to have a biological basis? What could it mean for something not to have a biological basis? What does this all mean for the relationship between biology as a discipline and its sister disciplines? A third problem that concerns me is interactionism itself. How does it manage to be virtually universally adopted and thus to lend itself to such radically different approaches? The suspicion is that it has become conceptually vacuous while acquiring the symbolic value of a membership badge, to be flashed upon entry into serious discussion: yes, I belong to the company of reasonable people; now let's talk about the real stuff. Though interactionism is not as innocent of substance as sometimes seems to be the case, and though, in fact, its multiplicity of meanings becomes clear when it is examined closely (see, for example, exchanges between Kitchener, 1978, 1980; and Lerner, 1980), one of the reasons it serves as an all-purpose slogan is that its users often agree on more than they realize. That set of shared assumptions, of course, about biological nature that can interact with environmental nurture is what occupies much of my attention in the pages that follow.
Though a full-scale critique of interactionism will not be undertaken here, psychologists and biologists continue to explore its many faces and to debate the relative merits of each. One meaning of interaction is statistical; it refers to the interaction between independent variables in an analysis of variance. When such statistical interaction is observed, an independent variable's effects on the dependent variable cannot be accurately described without reference to another independent variable. Though it has often led to senseless attempts to partition causal primacy according to the size of interaction terms, this kind of interaction is important to the view presented in the following chapters insofar as it directs attention to the mutual dependence of effective causes. (The importance of this principle does not, however, require the discovery of statistical interaction in any particular investigation. This is because results reflect the conditions chosen for that research; they may or may not produce interaction terms.) When the effects of a stimulus depend on genotype (actually, on the phenotype) or when the effects of a gene depend on the cell's location in an organism, the relativity of cause-effect relations to the context of investigation is crucial.
Another sense of interaction as I see it emphasizes the necessity of viewing transactions between an entity and its surround as aspects of a single system. When we speak of organisms altering their environments, both animate and inanimate, and being altered by their interactions, it is this mobile interchange that is being highlighted.
Hyland suggests that "interaction" has too often been treated as an answer or an explanation rather than the statement of a problem for which appropriate methods of investigation must be found. He was referring to the persons-situations debate in personality theory, which I discuss in the next chapter, but his description can also be applied, for example, to some research in behavior genetics. If analyses of variance are solutions in search of a problem, he concludes, one could also say that "the solution is the problem" (1984, p. 325; see also the commentary following Hyland's article).
In the next chapters, three fundamental phenomena of life processes will be reviewed: constancy, change, and variability. Each has been explained, completely or in part, by "nature" manifested in the genes, and each, when examined closely, reveals crucial aspects of ontogenetic processes. In Chapter 2, constancy in living things will be discussed, and so-called genetic constraints will be seen to be a function of phenotypic structure and function and of developmental context. The mutual dependence of effective causes, or reciprocal selectivity, will be described as basic to the understanding of biological forms, which are viewed as results of interactive construction, not manifestations of a preexisting plan. A consideration of change, in Chapter 3, will allow examination of the development and functional contingency of information and clarification of the way reciprocal selectivity works in ontogeny. Three kinds of variability will then be treated in Chapter 4, and the distinction between inherited and acquired characters (information, programs), on which natural selection has been said to rest, will be seen to be spurious, not only with respect to the description of individuals, but with respect to evolutionary change as well. In its place the notion of developmental systems will be advanced. This idea easily accommodates multiple developmental pathways in a way that the older idea of the inherited cannot (and constancy of pathways in a way that older concepts of the acquired cannot), allowing a consistent description of ontogeny, phylogeny, and the relations between them.
(Continues...)
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