Epigenesis 1. Inateness 2. Four Types of Questions (Tinbergen) 3. Epigenesis

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1 Epigenesis 1. Inateness A. Developmental Fixity B. The Strategy of Backward Causal Inference and Its Problems 2. Four Types of Questions (Tinbergen) A. Proximate B. Phylogenetic B. Ontogenetic D. Function: Survival Value (or Adaptive Value) 3. Epigenesis A. Umwelt 1. Assemble 2. Alter 3. Transduce 4. Modulate B. Umwelten and Complexity C. Generative Entrenchment D. Inateness Again

2 Page 2 1. Inateness A. Semantic Issue (i.e., what is meant by development): Developmental Fixity Developmental Fixity: Behavioral development in humans and other animals is fixed. Whatever the pattern of development, it is presumed to occur in the absence of any significant environmental input. Biologically, the most extreme form of developmental fixity is preformationism. Regnier de Graaf ( ) claimed that he could see a homunculus inside sperm. On this view, development is just a matter of growth and the role of the woman is to aid that growth. Regnier de Graaf From a cognitive point of view, the developmental fixity view can be traced back Plato s dialogs in the Meno (which means stable, fixed, 380 BC). In which Socrates uses a priori arguments for the view that some knowledge is innate.

3 Page 3 From a more modern perspective, the ideas of closed or open (with critical periods) developmental programs also have the idea of developmental fixity in it. And those who believe in developmental fixity turn to genetics for support. But, As we have already seen, the concept of a program is not fully adequate to explain development, even that of an open program, and Most importantly, there are no examples of developmental fixity (though towards the end of this lecture, I will talk about a reason why some features of early development may be more stable than later developmental stages, but this will have nothing to do with developmental fixity). The idea that developmental fixity is a consequence of genetics is a mistake. B. The Strategy of Backward Functional Inference and Its Problems Michel and Moore raise the important issue that how a scientist looks at a phenomenon (scientific perspectives as we talked about earlier), influences what kinds of questions will be asked and what causes will be looked for in development. From a functional perspective (in the evolutionary adaptive sense), a scientist will typically focus on the adaptive consequences of behavior and look for the sequence of developmental events leading up to the behavior in question. This is the strategy of backward functional inference. It is not an invalid form of inference, but it is easy to oversimplify development and view it as a predetermined sequence of events or stages. A scientist working from an ontogenetic perspective views development as a causal process in which a myriad of environmental, genetic, historical, and social factors play a causal role at each stage of development.

4 Page 4 Thus, a behavior may appear to be innate in the sense of being predetermined or developmentally fixed, when in fact it depends on specific environmental factors being present in its ontogenetic environment. Laughing Gull Chicks: Laughing gull chicks will approach and peck a parent s beak as a consequence of a croon call by a parent. Parents then regurgitate food for the chick to eat. From an adaptive functional perspective, failure of a chick to approach would be very maladaptive! 1. From an adaptive point of view, one might conclude that laughing gull chicks have been designed by natural selection to approach in response to a croon call, which in turn suggests that the approach response to croon calls is genetically programmed. 2. The adaptive design view is further supported by the fact that laughing gull embryos can apparently discriminate croon calls from other adult calls. 3. Thus, because approach to croon calls is adaptively important AND it is observed early in development, it appears to be innate in a predetermined and genetic sense sense (i.e., part of a closed program). From an ontogenetic perspective, however, the focus must be on the causal conditions that precede the behavior. Impekoven and Gold (IMPEKOVEN, M., AND P. S. GOLD Prenatal origins of parent-young interactions in birds: a naturalistic approach. Pp in Behavioral embryology (G. Gottlieb, Ed.). New York, Academic Press.) showed that the ability of gull chicks to respond appropriately to the croon calls of their parents depends on the chicks having heard the calls before they are hatched. Here again we see the importance of social levels of organization. Parents when incubating the nest make croon calls. Several days before the chicks hatch, they begin to peep, which stimulates the parents to croon even

5 Page 5 more. Chicks raised in incubators do not have the typical approach response to croon calls, but instead crouch and hide. From an ontogenetic perspective croon calls emerge from a myriad of causal conditions and are not predetermined to occur regardless of conditions. This is compatible with the behavior s evolutionary adaptive function, we just need to keep in mind that adaptive behaviors need not be designed in early development or predetermined. 2. Four Types of Questions (Tinbergen) Tinbergen (Tinbergen, N. (1963) On Aims and Methods in Ethology. Zeitschrift fur Tierpsychologie, 20(4), ) emphasized that many confusions can arise in the study of the development of behavior if different questions or perspectives are not distinguished. Tinbergen distinguished four such questions: mechanism, survival value, evolution (phylogeny), and ontogeny. A. Mechanism Questions of this type are about the mechanisms that cause behavior (e.g., the influence of hormonal and neural events on behavior or the stimuli that evoke responses in animals). Crooning in laughing gull chicks, chemical signals given off by newly emerged works in army ant colonies, increased estrogen levels in ringdove females in response to male courtship behaviors. B. Phylogenetic Questions of this type are the longer time scale of causal conditions that affect behavior over evolutionary time. Time scale is over many generations. In one sense, phylogenetic causation is proximate causation operating on a much larger time scale. For example, if fall webworms that twitch synchronously when flies or wasps land on their web have a slightly better chance of surviving, then in the long run these repeated proximate causal interactions will ultimately result in twitching as an adaptation because twitching worms will come to make up the population.

6 Page 6 Behavior leaves few cues in the fossil record, so it is difficult to reconstruct the causal conditions of current behavior. Sometimes we can use phylogenetic analysis of other characters (including genetic relationships) to suggest plausible precursors for presently observed behaviors. For example, ritualized courtship behaviors (e.g., in mallard ducks, ring doves, etc.) may have evolved in part from stereotyped preening behavior (exaptations). Finally, although phylogenetic causation focuses on the historicalevolutionary relationships between behaviors, all the causes we have discussed occur on a smaller time scale within phylogenetic causation. C. Ontogenetic Ontogenetic causation concerns the time span of an individual. It focuses on the causal conditions preceding each stage of development, and thus can also be viewed as proximate causation on a larger time scale as with phylogenetic causation, however, all four causes are playing a role. D. Function: Survival Value (or Adaptive Value) 3. Epigenesis Questions concerning survivability and fecundity. Finding a mate. Selecting a mate. Holding territories and resources. All of the previous causes have implications for survival value. A perspective on ontogeny where development is (i) not predetermined or fixed, (ii) not reducible to genetic programs, (iii) a process involving interactions from many levels of organization acting on each stage of development,

7 (iv) an active process of unfolding and emergence in which developing organisms, assemble, alter, transduce, and modulate their environments. A. Umwelt Von Uexküll concept of Umwelt is the notion that the environment of an organism consists of those features it perceives and reacts to. Page 7 The Umwelt of a tick and their detection of butyric acid given off by mammals. Developmentally, the Umwelt of an organism changes as the organism develops. Newborn rat pups are born blind and deaf with limited sensorimotor capacities, and as sensorimotor abilities emerge, so does their Umwelt.

8 Richard Lewontin further elaborated the active role of the organism in defining its Umwelt. Page 8 1. Assemble: Organisms partially assemble their environments, e.g., many birds build nests, social insects build nests, spiders weaving webs. 2. Alter: In interacting with their environments they alter them. This is not necessarily the same as assembling. For example, masses of locusts devouring vegetative matter in their path or elephants pulling down trees to feed on the leafy branches. 3. Transduce: The take in energy and perceive stimuli and produce physical products and behavioral interactions. Consider a female mammal nursing her young. She takes in energy in the form of food and transforms it into milk. Stimulation from her offspring induce the release of oxytocin, which causes milk letdowns. 4. Modulate: They regulate their environments. Even Army ants can control the temperature and climate of their nests. Epigenesis is thus a view that development is not a passive unfolding of stages of development, but an active process in which organisms create (partially) the environments in which they develops. B. Complexity of Umwelten Recognizing differences in the Umwelten of animals reveals both differences in complexity across taxa and during stages of development. This has important implications for experimental research in developmental psychobiology: the more complex the Umwelt of an organism the more likely it is that: Biases of oversimplification can creep in. This can happen for both innate and acquired research. (e.g., intentionally in new born humans, or Skinner boxes and learning).

9 Page 9 The more difficult it is to generalize results from the laboratory to the more complex Umwelten of organisms in their natural environments. But: the more we understand an organism s Umwelt, the better able we are to investigate it in the laboratory and generalize results we discover. It may be a good strategy to start by studying organisms with simpler Umwelten first, e.g., ticks or organisms with more complex Umwelten early in development, when their Umwelten are simpler. C. Generative Entrenchment Epigenesis is a generative process meaning that each stage of morphological and behavioral development is causally generated from previous stages and the environment (in particular the Umwelten of the current and previous stages). William Wimsatt first noticed changes early in development (e.g., genetic mutations, changes in the environment), will often have more effects on development than changes later in development, because early changes can causally cascade through development. Now, if most arbitrary or random changes in development are deleterious to the adaptive function of the developing organism at any later stage of development, then all things being equal, early stages of development will tend to be more stable and the interactions with the environment will be more stable. This explains von Baer s (a 19th century German developmental biologist) who argued that Ernst Haeckel s view that ontogeny recapitulates phylogeny is not factually correct; embryos (e.g., across vertebrate taxa) tend to be more similar early in development and diverge later.

10 Page 10 Developmental Stages Environmental Influences Causation Developmental Time

11 Page 11 Ear ly Change Late Change D. Inateness Again If we mean by innate behavior, behavior that emerges early and is critical for later development, then the more generatively entrenched a behavior is, the more innate it is. Notice that innateness is no longer tied to genes but rather to a component s (i.e., a gene, environmental factor, anatomical or physiological structure or process) generative role in development. Innate factors will tend to occur early in development and resistant to evolutionary change.

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