Levels of Organization and Causation 1) Levels of Organization a) Composition b) Interactions 2) Time Scale a) Composition b) Interactions 3) Scientific Perspectives and Organization 4) Causation a) Aristotle’s Four Causes i) Material Cause ii) Efficient Cause iii) Formal Cause iv) Final Cause b) A Reformulation of Aristotle’s Four Causes i) Material Cause => Composition ii) Efficient Cause => Proximate Cause iii) Formal Cause => Organization iv) Final Cause => Function (1) System Function (2) Evolutionary adaptive function 5) Army Ants page 2 1.A. Levels: Composition Video: Fall webworms One meaning of the term Levels is composition: A thing X is at a higher level of organization than a thing Y if X is composed of Ys (and possibly other things). For example, cells are at a higher level of organization then DNA molecules, because among other subcellular structures, cells are composed of DNA. Galaxies Planets Ecosystems Societies Groups, Populations Multicellular Organisms Cells Subcellular Structures Proteins, DNA Subatomic Particles Notes: page 3 Compositional Levels do not reveal how the components at one level interact to produce the behavior and organization of things at a higher level. Thus, for example, merely knowing that a group is composed of individuals of a particular species (e.g., army ants, wild dogs, or humans), tells us very little about the social structure of the group or how the individuals develop in the context of the group. 1.B. Levels: Interactions Another meaning of Levels is the number, strength, and kinds of interactions among components at different levels of composition. The interactions of components at a given level constitute levels of organization. Organization often emerge from simple interactions of components as we saw with fall webworms and as we will see with army ants below. Example: Aggregation Rules Complex systems, however, are not limited to interactions (i.e., interactional complexity) with components at one level of compositional. For example, oxytocin, is a peptide hormones, involved in the process of birth and nursing in mammals and social behavior, but there are many other physiological processes and behavioral interactions required for these processes to occur. The particular function of a hormone such as oxytocin on the state and environment of an organism, i.e., its context or the situation. Vasopressin, a peptide hormone closely related to oxytocin is best known for regulating kidney function. If the vasopressin gene is defective, mammals such as rats will develop diabetes insipidus. But, it also plays a role in social behavior and anxiety and appears to modulate early locomotor and social development in rats. It may also modulate early sensorimotor development, making it a prime candidate for playing a role in autism. The figure above is a rather rough ordering of different systems in terms of the levels of complexity of their interactions. Note, for example, that although galaxies are at a very high level of composition (i.e., they are composed of many different kinds of things) they are at a low of interactional complexity. That is, many of the things that a galaxy is composed of do not directly interact other than overall gravitational forces. For example planets and any living things that may be Notes: page 4 on them have negligible influences on the organization of galaxies—of course galaxies are composed of complex systems, it is just that they are not complex systems by virtue of being composed of complex systems. Groups, Populations Multicellular Organisms Societies Proteins, DNA Subatomic Particles Galaxies Planets Cells Subcellular Structures Simple Ecosystems Complex Interactions A. Composition ! Interactions If we plot systems in terms of their compositional level versus their interactional level, we see that there is not a simple relationship between the two. Confusing these two levels can and has led to oversimplifications of the relationships between behavior and development, genes and development, and evolution and development. Later we will see that these two levels have different meanings of Causation associated with them. Notes: page 5 Composition Galaxies Planets Ecosystems Societies Groups, Populations Multicellular Organisms Cells Subcellular Structures Proteins, DNA Subatomic Particles Interactions Notes: page 6 2.A. Time Scale ! Composition When we look at the time scale of processes, we see that compositional level and time scale are closely correlated (the figure below is only an approximate representation of the relationship). Composition Galaxies Planets Ecosystems Societies Groups, Populations Multicellular Organisms Cells Subcellular Structures Proteins, DNA Subatomic Particles Time Scale Notes: page 7 2.B. Time Scale: Interactions Interactions As I mentioned last time, a myth of developmental psychobiology is that if systems (e.g., the development of an organism, ecological systems, species) change on different time scales, then one can study one type of system relatively independently of the other. This can lead to error, as we will see in this course. Notes: page 8 3. Scientific!! Perspectives and Organization Cricket userdict /mypsb currentpoint /newXScale /newYScale {}Software store /psb /mdnewHeight newWidth known{/CricketAdjust /picOriginY /newHeight pop /pse store /newWidth def {} store /mypse 427 493 exch div div /pse exch picOriginY def def defload picOriginX /picOriginX truedef def}{/CricketAdjust sub sub def exchpop def def false def}ifelse /psb /pse /mypse /mypsb load load store Earlier I said that the function of a hormone such oxytocin depended on the state of an organism (e.g., whether it is pregnant) and its environment (including social context). From a researcher’s point of view, questions of function and mechanism Notes: page 9 lead to different views or perspectives of a system. Depending on how we view a system can lead to different ways of decomposing a system that is at the same level of composition. In the diagram above, I have illustrated different perspectives one may take in studying a piece of granite and a fruitfully. In the case of the piece of granite, taking different perspectives leaves us with the same, relatively simple description of components. However, with the fruitfully, depending on the perspective and questions asked, we get descriptions of complex systems, which differ according to perspective. 4. Causation By our ordinary meaning of causation, we understand the relationship between cause and effect as one of conditions that bring about certain effects. For example, throwing a ball and thereby breaking a window. The summation and integration of excitatory and inhibitory postsynaptic potentials bring about action potential in a neuron. Causation in this sense is proximate, which means that the conditions bringing about an effect are near their effects in space, time, and typically at the same level of organization. Aristotle, a famous ancient Greek philosopher, had a more general notion of causation. He thought of causes as reasons or factors explaining the objects and processes that exist in the world. For Aristotle there were four basic causes: 1. Material Cause: The material of which a thing is made. 2. Efficient Cause: The conditions that combine to produce an effect from a cause. 3. Formal Cause: The shape, configuration or type of thing something is. 4. Final Cause: The purpose or end of a thing or process. Notes: page 10 Aristotle held that anything could be explained in terms of these four causes. For example, the material cause of a statue is the stone it is made of. Its efficient cause is all the chiseling required to transform the stone into a shape or configuration, its formal cause, which was the purpose or aim of the sculptor who produced it, its final cause. There is a ring of truth to Aristotle’s four causes as explanations for why a thing is the way it is, but Aristotles four causes require a modern reinterpretation in light of modern science. Indeed, there are modern ideas that correspond neatly to Aristotle’s four causes. They are: 1. Composition: Composition corresponds very closely to Aristotle’s notion of material cause. It is the components, entities and processes that compose a system at some level of organization. For example, DNA, RNA, and Proteins are among the components making up cells. 2. Proximate Cause: Proximate cause also corresponds very closely to Aristotle’s notion of efficient cause. These are the causal conditions specified in mechanisms for how things work, such as the firing of a neuron, or hormonal factors that modulate reproduction and maternal care of young. 3. Organization: Is one aspect of Aristotle’s notion of formal cause, namely the idea of configuration. But, specifically we are primarily concerned with the configuration of interactions. Interactions connect organization to composition and proximate cause. A. Interactions are among components of a systems, and B. Interactions are proximate causal relationships among components. 4. Function: The notion of function is the biggest departure from Aristotle’s system. Final causes assume a purpose for a component or system. This is fine for ordinary human artifacts. A purpose of a table is to set things on, and this is a reason for it existence, but final causes are neither testable nor explanations of natural systems. Thus, in biology and psychology we replace Aristotle’s notion Notes: page 11 of final cause with two related concepts of function. Functions in either sense are understood in terms of their consequences. A. System function: This is the role of a component in a system. For example, a function of a neuron is to modulate action potentials. A function the heart is to circulate blood. System functions depend on the composition of components, their proximate causal interactions, and how these interactions are organized (e.g., the configuration of excitatory and inhibitory inputs into a neuron, branching nature of the circulatory system). B. Evolutionary adaptive function: This is the role of a component in the survival and reproduction of a system in a larger ecosystem or environment. Typically, adaptive functions are ascribed to components and characteristic of individuals, but they may apply to entities at other levels of organization such as groups, ecosystems, and species. Army Ants Army ants are an interesting example of the complexities of organization and the different kinds of causes operating during development. What makes this example especially interesting is that individual insects such as ants are in many respects much “simpler” (this is a relative term, ants are nevertheless quite complex systems) than birds or mammals in the complexity of their nervous systems, their behavioral repertoires, and in their capacity to learn. T. C. Schneirla studied army ants (Eciton hamatum and E. burchelli; other websites 1, 2, 3) throughout the 1940s. Army ants are nomadic foraging social insects. Unlike many other social ants, wasps and bees they do not build permanent nests. [Videos: 1 2 3 4] One of the more remarkable aspects of army ant social behavior is that individual development is highly coupled with social behavior and individual development drives social behavior. Notes: page 12 Cycles of Behavior Army ants (50,000-120,000 in a colony) exhibit daily or diurnal cycles of functional activity. Functional activity includes daytime foraging (when light, temperature and humidity are best), thermoregulation and climate control in the bivouac. These functional cycles are largely under external control of the daily light cycle, temperature and humidity conditions. These are external Zeitgebers. They go through Nomadic and Statary phases of colony behavior that are tightly coupled with development of offspring. Interestingly, these cycles are not driven by external Zeitgebers but emerge from the individual interactions of workers, new workers and pupae. Notice that all four types of causes are present Schneirla’s representation. ! The Colony Functional Cycle refers to the functions performed by the colony such as foraging and incubation, care and feeding of the next generation. ! The Brood Cycle corresponds approximately to the notion of proximate causation, though later we will distinguish types of proximate causal relations that occur at different time scales such as ontogenetic causation. The Typical Raiding Pattern illustrates one aspect of the organization in army ants, particularly, foraging search patterns, which are the result of simple individual rules of behavior. Notes: page 13 Notes: page 14 How are these functional cycles, and synchronous reproduction and development generated? According to Schneirla they are due to local interactions between newly emerged workers and the current work force and subsequently reciprocal interactions between workers and the developing brood. 1. Newly eclosed brood (first spike above threshold): Newly emerged works increase the activity levels of old workers in the workers lick and groom the newly emerged workers due to chemical signals. Notes: page 15 2. Developing larvae (the second more gradual rise): Workers are attracted in the presence of larva pheromones and nutritive secretions that the workers ingest. Larva also become more active in the presence of workers, who feed them. Activation of workers spreads throughout the colony. This process of behavioral excitation and activation of workers is not merely timed by the length of time it takes the larva to develop. Increase activation stimulates the nomadic phase, foraging activity is increased, increasing the food supply needed for growth and maturation of larvae. Notes: page 16 The contents of Psychology 113, Developmental Psychobiology, as taught by Dr. Jeffrey C. Schank during the winter 2012 academic quarter at the University of California, Davis, are COPYRIGHT Jeffrey C. Schank (c) 2000-2012. All federal and state copyrights are reserved for all original material presented in this course through any medium, including lecture or print. Notes: