The organism of behavior

December, 2020 The organism of behavior A.P. Rabanal † Behavior analysis relies on the idea of a science dealing with specific interactions between the organism and its environment. Yet it is not always clearly defined what does “organism” mean. It does not even seem to be an agreement on it within biologists beyond the idea of a physically sustained and self-maintaining entity (Ruiz-Mirazo, Etxeberria, Moreno and Ibáñez, 2000). While a technical and foundational concept of “organism” may be sought in biology, whether or not it is of psychological interest is another issue. This is not to deny species-intrinsic traits nor the role of neurobiological mechanisms, but to stress that adaptative behavior is best exemplified in situationally-bounded organisms. Within the behavioral community positions vary from those who accept organismal variables (e.g. Donahoe and Palmer, 2004) to those who propose not considering them at all (e.g. Roche and Barnes, 1997). I will contend that disagreements arise from the roles the organism is said to have in behavioral settings. I will also argue for the organismal level as appropriate for explaining behavior in an attempt to reconcile physiology with behavior analysis through a selectionist approach. Levels and Organization Living systems are hierarchically structured in a set of successive organization levels from the genome to the individual (Salomon, Berg and Martin, 2008) with elements belonging to intermediate levels between genes and cells (e.g. proteomes or transcriptomes) or between populations and ecosystems (e.g. communities) (Pavé, 2006). Biological complexity is one of downward organization and should be explained in terms of both reduction and integration. Bunge and Mahner (1997) propose the following composition criteria of biological levels: L1 = Elemental cellular level def= set of every elemental cell L2 = Compound cellular level def= set of every compound cell L3 = Organal level def= set of every (multicellular) organ L4 = Multicellular organismic level def= set of every multicellular organism L5 = Population level def= set of every biopopulation L6 = Community level def= set of every community (or biocenosis) L7 = Biosphere level def= set of every biosphere 1 December, 2020 Non-living elements encompass oxygen, water, soil, or minerals with factors to consider such as spatial (resource distribution or physical geography) and temporal restraints (circadian rhythms, lunar cycles, or seasonal changes). MacMahon, Phillips, Robinson and Schimpf (1978) indicate that biological complexity culminates with the organism as an ecological unit as levels not containing living systems are not properly biological. Levels can be then taken as clusters: L1 = Cellular level L2 = Organismal level L3 = Population level L4 = Community level L5 = Ecosystem level Levels are sets and hence conceptual in nature. Conversely their components are collections of living things from cells up to organs. We can further complement this classification with a functional criterion. As basic units of life, cells are characterized by their morphology and metabolism. Ensembled or associated cells with extracellular macromolecules form compounds which perform specific functions. A complete organ is a group of tissues with a same function (i.e. a parenchyma) and a supportive stroma. Tissues and organs make up organ systems which accomplish complex coordinated functions such as breathing, digesting, protecting the individual, or processing information. The organism is thus a functional individual supported by subsystems operating together. Organisms can also be regarded as the highest unit of life as they are not proper subsystems of other (Bunge and Mahner, 1997, p. 149). They are instead constrained by contingencies to survive. Two or more organisms of the same species confined in space and time with demographic parameters constitute a population where intraspecific interactions as cooperation or competition arise (Pavé, 2006). Any system containing at least one organism provided with the appropriate nutrients can nonetheless be an ecological system by its own (e.g. the D. audaxviator specimen) (Strickland, 2008). A community involves different species populating the same area and where symbiotic interactions such as competition, predation, commensalism, mutualism, and parasitism occur. Lastly, an ecosystem contains nonliving elements and is spatially limited leading to the development of niches, immigration, emigration, or trophic dynamics. Whereas cells communicate by signaling through direct contact or via a local regulator, chemical compounds (e.g. neurotransmitters) or hormones in the blood torrent, for the organism communication is rather a transaction of energy with the external environment. The source of energy that powers most life such as electromagnetic waves can be traced 2 December, 2020 in the sun. Although cells capture, transfer, store and use energy, they do not produce it. Most actions of an organism involve a series of energy transformations as kinetic energy is converted to potential energy (Salomon, Berg, and Martin, 2008, p. 152). Heterotroph organisms seek food sources and potential mates in their immediate surroundings. They are open systems exchanging energy and releasing it by engaging in various activities. The organism is the basic ecological unit along with communities but there is also a qualitative break between them (Pavé, 2006, p. 47). Species share the highest level of genetic similarities transmitted to their offspring while the community is genetically heterogeneous. Biocenosis or communities are not living entities but rather a collection of biosystems from prokaryotic cells (e.g. archeas or bacteria) to morphologically separated individuals with joint metabolism (e.g. corals). Neither internal parasites nor certain algae are proper parts of their hosts as in the case of lichens until each cyanobacterium separates from their fungal filaments. A spatial restraining is ultimately imposed whether it is a lipidic membrane or epidermal tissue as individuals selectively interact with their environments. “Primitive organisms” may have evolved from replicator molecules by satisfying the requirements of relative independence, division from an external environment, and confinement of biochemical activity. There are thus at least two meaning of “interaction”: one involving inter- or intraspecific relationships affecting reproductive success; and another one involving the transference of energy between the organism and its environment with the most proximate components forming strongest interactions (MacMachon et al., 1978, p. 702). Yet Pavé (2006, p. 46) suggests that basic interaction processes can be found in any level. Reproduction, induced degradation, competition, association, and adaptation are analog processes to be traced from the cellular level. For instance, protein degradation can be compared to predation; and metabolic cooperation to altruistic behavior. What matters is whether organisms have emergent properties not attributable to their components. For example, most body cells are replaced through development, but the individual remains relatively the same once an adult. Lidicker (2007) enumerates some properties related to the organism such as a coordinated trajectory, homeostasis, resources acquisition, mate finding, or defense against potential menaces. Demographic density, social organization, sex ratio, speciation, genetic heterogeneity, or evolutionary arms race are instead populational constructs. We have to consider then to what extent behavior is an emergent function of the organism as a whole not belonging to another level of organization. Organisms and Environments Compared with the idea of the organism as an assemblage or “vehicle” of replicators (Dawkins, 1986), or the rather eccentric depiction of the individual as a “host” of responses (Baer, 1976), the notion of the organism in psychology is usually confined to that of a “selfcontained biologically bounded individual” that may impose or not constraints on learning (Roche and Barnes, 1997). It has even been said that there can be no complete account of behavior without considering some biological characteristics of the individual 3 December, 2020 (Kanfer and Phillips, 1970). But psychological statements are usually regarded as concerning learning contingencies without the need of further specifying a role for the organism. Individuals are of most importance from early behavioral ecology (Moen, 1973) to current trends (Owens, 2006). One can set aside some of these topics as concerning biologists insofar ecology studies the interactions of organisms and environments across all ecosystems (Malmstrom, 2010). It happens that this is claimed to be the subject matter of behavioral analysis as well. For instance, Woodworth (1958) defines psychology as the scientific study of the activities of the organism in relation to the environment. For this reason, Roche and Barnes (1997) disdain the loose organism-environment interaction concept by calling for targeting only “special types of behavior”. But it would be easier to consider that some approaches overlap with other disciplines just tackling distinctive aspects of behavior. Behavior analysis is a part of biology as responses refer to processes exemplified by living organisms (Donahoe, 1996). It is no more dependent on physiology than physiology is on cellular biology, or cellular biology on genetics. Yet it is neither less interdependent on them than they are on each other. And although controversies remain whether endosymbionts or insect colonies are organisms (see Pepper and Herron, 2008), there are no major controversies in psychological settings as subjects of research are easily identified as separate higher vertebrates. Nevertheless the organism can still be considered no more than a container providing external protection for the organs. Body systems perform complex functions such as breathing, digesting, excreting, or regulating bodily functions. Sessile organisms as mussels can travel via water currents or have a motile phase during their development as sponges. Tropisms and nastic movements are to be distinguished from kineses and taxes which are orienting reflexes of motile organisms and dependent upon current environments. It seems that organisms only move bringing everything inside with them as once reached fetal viability the ability of the individual to survive depends thereon its success in coping with nature outside the uterus. From this follows that an “organism” is an individual that embraces the highest level of organization of living forms. But it also keeps a functional control on the exchange of matter and energy between the environment without which their internal milieu would not survive. The exchange of gases occurs in cellular membranes and some organisms must breathe in oxygen available in the environment. While metabolic reactions are responsible for converting chemical energy to ATP, the energy itself is derived from nutrients from external sources. Except for binary fission, budding, or sporulation, reproductive success cannot be made without finding a proper mate. Sensorial deprive can also be detrimental to individuals. Individuation processes instead encompass the success of reproduction and development (Ruiz-Mirazo et al., 2000). The notion of “organism” may be then looked up in its interaction with its surroundings to increase its survival rate. The skin actually entails a pivotal role as there are no 4 December, 2020 functional relationships if environmental events or stimuli never reach the individual (Moen, 1973, p. 18). But neither are if no appropriate input and output channels are considered. Even Baer (1976) suggests that once the organism enters the operant chamber, the concern is explaining behavior under contextual control with only a “single parametric specification of the health or the organism” (p. 89). Palmer (2004) recommends an analysis of transdermal processes (though and across the skin) as complementary phases. Engaging with changing environments requires learning novel responses and natural selection has provided us with a genetic endowment to reliably respond to relatively stable biotopes as evolutionary strategies. This is criticized by Bunge and Ardila (1987) for outputs are a function of both stimuli and inner states. The same output can be expected only by monitoring the state of the animal in each trial. Any transaction with an environment can be represented as O = M x I where O refers to the output as a product of both the input I and the summarized properties M of the organism (Bunge, 1975, p. 63). When M is nowhere stated, the model cannot explain outcomes in terms of mechanisms but only control variation resorting to M = O x I-1. Yet an interpretation of the role of organismal variables (internal states) would remain absent. However not only physiological mechanisms (e.g. synaptic efficacies) are needed, but also functional explanations: A S C R C S Figure 1. Stimuli have been dubbed respectively as antecedent and consequent while the operant class has been changed to a generic response class. Even if internal mechanisms supplement this account with mediating processes, they do not substitute the need for specifying environmental contingencies. Adapted from Zilio (2020). Whether psychologists study the behavior of plants, flatworms, or computers; or if the choice of the organism is merely incidental for facilitating experimental purposes raises the issue of the idea of a “psychological organism”. Is the decision made upon ethical restraints or the easiness in the handling of the animal? Or does the choice of a specific organism add something to the experiment? The vast majority of animals are of no interest for the psychologist but only those capable of learning. Therefore, only animals endowed with a nervous system that enables them to react to their environments and learn upon their consequences can be classified as subjects of interest (Bunge and Ardila, 1987, p. 29). Until we have more insight on the adaptation and brain chemistry of other species (e.g. reptiles and fishes) one may rely mainly on mammals and birds. Matthews (1985) claims 5 December, 2020 that what accredits a “psychological organism” is a minimal plasticity which enable it to learn. Although “psychological events” are more than biological events, they are no less than them. The “psychological subject” is a “biological organism” that engages in specific environmental relations whether the contiguity and discrepancy requirements are met. A morphological account of the organism would take the individual as a point in space (Palmer, 2004). Psychology needs instead a functional account of the organismenvironment system as one cannot separate both as long as responses and stimuli are codefined. The cell is morphologically (structurally) limited to the membrane, but functionally related to the extracellular space where ions and organic molecules will trespass the cytoplasm in order to ensure its survival. As skin, horns, carapaces, or scales functionally interact with the environment, this perspective encompasses both physiology and psychology as a feedback scheme through a behavioral (Fig. 1) or a biological approach (Fig. 2): A S O C C S R Figure 2. An expanded biobehavioral paradigm when organismal variables (O) and inner states are taken into account between antecedent stimuli and response classes, but not substituting the contingencies which remains either on the internal or external environment of the individual. It can be said that intraorganismal variables also learn to respond to external stimuli like the immune suppression with the administration of saccharine sodium associated with cytophospane. But the conditioning process is found in environmental contingencies. Even more, organisms differ in their biological structures shaped by natural selection but also in their behavior from one environment to another (Donahoe and Palmer, 2004, p. 18). Behavioral variation is retained by the ancestral environment but it is enriched following birth by the individual environment. Although we lack “behavioral fossils” such as changes of synaptic efficacies, we can find endocasts and permanent products made by our ancestors. Yet it is ultimately the environment that favored some responses over others due to selective pressures. Variability diminishes when an “action strategy” is learned, but when consequences that usually follow it fails, there is an increase of 6 December, 2020 variability as it would have returned to an exploratory phase (Veá, 1990). Organisms that learn to respond in appropriate ways to novel stimuli were more likely to leave offspring, so natural selection favored organisms with plastic neural systems. And if different operants are selected in similar environments, then the response repertoire is enlarged leading to adaptive behaviors of increasing complexity. It is not superfluous to say that learning is no more nor less a fundamental principle for adaptative behavior but a second kind of selection only behind natural selection (Skinner, 1981). The evolution of species and the individual development of the organism create the conditions upon which environmental contingencies operates thus shedding light on the roles of biological variation for explaining behavior. States and mechanisms Every individual is embedded in an environment so we have to be aware of both the biological characteristics of the organism and the physical surroundings where it develops and learns. Organismal variables that may influence the trajectory of behavior can be considered either: 1) internal states or biological conditions; or 2) intermediate mechanisms. Certain physiological processes such as blushing, perspiring, crying, or sweating are responses in themselves. As the organism is intermittently present in any behavioral event, only variables that facilitates or interferes response functions such as chromosomal alterations, early infections, age, nutritional state, neurological conditions, or physical impairments are to be considered (Kanfer and Phillips, 1970). It is to notice that hereditary factors and pre- and perinatal factors are themselves not mechanisms but variables that adjust the CNS of the organism. They alter responding or impose constraints on learning which remains a function of contingencies. Behavioral repertoires and learning histories are also considered organismal variables as are retained in the organism (e.g. through generalization of contextual cues via the strengthening of neural connection in the associative cortex). All these are internal states that correlate or interfere with the establishment of contingencies and thus are province of basic and applied psychophysiology. But organismal variables can also be considered mechanisms on par with environmental events. This is not to say that motor responses are explained alone by neuromuscular mechanisms, but rather that a response is explained by the environmental contingencies, that is, by the circumstances that set the occasion for the emission of a motor response. Physiological mechanisms mediate between the antecedent event and the consequent event manipulated as contingencies (Fig 2). This can be seen as the most prosaic proposal of reconciliation of behavior analysis and neurosciences (see also Zilio, 2013). A somewhat sophisticated approach is proposed by Donahoe and Palmer (2004). Complex behavior is said to emerge as the cumulative product of selective processes acting over time akin to biological evolution. Simple responses can lead to complex phenomena such as thinking or reminding due to the selecting effects of the individual environment. The experimental analysis of behavior and research on the underlying physiological 7 December, 2020 mechanisms support this thesis. Temporal contiguity can be traced in the intracellular events leading to LTP and the discrepancy requirement has its neural counterpart in the increase of dopamine release from midbrain cells if these were not currently activated (Donahoe, 2003, p. 10). The standard S-R scheme can be explained in terms of stimuli acting upon receptors to produce activity in the primary sensory cortex. The stimulation of the sensory and the motor association cortex ultimately led to activity in the primary motor cortex causing measurable responses at the behavioral level (Donahoe and Palmer, 2014, p. 12). The same proposal is exposed by Bunge and Ardila (1998): “Given a stimulus s acting on an animal b, b contains a neural system n controlling the performance of an overt response, in such a way that the latter is determined jointly by s and by the state of n at the time the stimulus (or its transduction) reaches n” (p. 169). A purely functional account of behavior is likely to be inviable without considering feedback mechanisms. An organism with a different neural structure will presumably still accomplish the S-R scheme, but the task will be restricted to prediction and control until we obtain information about its internal states. However, while the notion of organismal events playing a mechanistic role is present, understanding how the environment exerts discriminative control provides a complete account of adaptative behavior. Roles and Functions In multicellular animals, an immense number of regulatory mechanisms have evolved. All organs and tissues function for the maintenance of constant conditions in the internal environment (Hall, 2016, p. 4). Some of these major body functions comprises extracellular fluid transport, nutrients provision, removal of metabolic end products, protection, and reproduction. Of particular importance and directly related to behavior are the musculoskeletal system and the regulation of body functions by both the nervous and the endocrine system. The former provides the organism with motility and external protection while the latter detect energy changes in the environment, are involved in locomotion and regulate internal cellular functions. The notion of “biological function” can be traced in ethological concerns for proximate causes (mechanisms) and adaptation values albeit it has an ancient origin. (For a historical review of the term see Toepfer, 2011). It usually means the activity of a component of a system so it can be identified with the set of processes occurring in it (Bunge and Mahner, 1997, p. 155). Different functions may fulfill a same biological role (e.g. claws, jaws, or stingers for protection). Pumping blood, beating, and emitting cardiac sounds are specific functions of the heart in the cardiovascular system but supplying oxygen and nutrients through blood circulation are their role. Functions describe processes or activities rather than traits (i.e. inherent properties) and are enacted by a biological system at an emergent level. (See also Farnsworth, Albantakis and Caruso, 2017). Behavioral responses are functions performed by an organism within 8 December, 2020 the context of a population, a community, or an ecosystem. Social behavior and interspecific activities are displayed in populational and communitarian dynamics. As every organism is a member of a larger ecosystem, it can also interact with its physical environment (e.g. great apes foraging techniques from antlers or rocks which can play a role for hunting). Thomas (2017, p. 16) opposes causal roles to selection functions so explanation of a biological function is to be found within its evolutionary history. This does not deny that functions of a system are identified with the processes occurring within. It simply expands the focus on whether such activity played a significant role during evolution, or whether it was a byproduct, or perhaps a spandrel. Selected effects favor certain processes as mechanisms over others for increasing inclusive fitness or reproductive success so are of special interest of evolutionary biologists while other accompanying effects may be of interest for the physiologist and the physician. It is the case of the function of a component of the visual system such as the eye. It may have contributed to the general fitness of an organism by enhancing light detection. But the functions of their components (cornea, sclera, pupil, iris, lens or retina) are the many activities they perform (light refraction, dilatation, accommodation, aberrations decrease). A minor refractive error may have no played a significant evolutionary role but is certainly of concern for the ophthalmologist. Structures with no apparent function (e.g. deer antlers) may play a significant role (e.g. sexual selection or protection from the environment). Conversely, the appendix carries out immunological activities, but its role is apparently insignificant (Bunge and Mahner, 1997, p. 156). Furthermore, functions of a trait are intrinsic properties of individuals, but roles are related to the part they play in larger environments (e.g. payoffs, kin selection, or prey adaptation). For avoiding confusion with its current usage, one can designate the “biological role” as the evolutionary or physiological functions of a biosystem whereas “function” will simply mean an activity that is carried out without raising the issue of its evolutionary origin but what it does and how it operates (Thomas, 2017, p. 17). In matters of explanation, behavioral analysts are concerned with ultimate and proximate causation whereas the former is the domain of both evolutionary biology (phylogeny) and developmental psychology (ontogeny) and the latter involves the discovery of mechanisms as a domain of physiology and behavioral neuroscience. Yet changes in the immediate environment alter synaptic efficacies that in turn influence behavior. Both environmental stimuli and neural mechanisms are deemed as proximate causes for learned behavior (Schlinger, 2015, p. 2). This leads us back to Skinner (1981) proposing that selection by consequences parallels natural selection by shaping and maintaining behavior not in the species, but in the organism. Behavior analysis addresses learning and adaptative behavior with respect to environmental contingencies while physiology accounts for the biological processes that underlie functional environmental-behavior relationships. 9 December, 2020 The organism as an open system interacts with its external environment and is subject to external and internal disturbances. Although cells interact with the outside thanks to their semipermeable membranes, the organism is the last living element in the process of energy conversion as it is not a proper part of any emergent system. An individual endowed with a physiological machinery constantly engages with a changing environment settling the occasion for natural selection to occur. Going back to systems physiology, one can advert that all processes of the organism usually refer to the internal milieu as if it were auto-efficacious (i.e. without an explicit reference to external cues, trigger signals, or environmental stimuli). It seems like a matter of common sense that olfactory receptors cannot transduce any chemical information if there is no object to be perceived except for olfactory hallucinations due to temporal or orbitofrontal lesions. The nervous system has evolved to meet the demands of interacting with and adapting to the environment with specific cortical areas (e.g. the frontal association cortex) largely programmed by learning experiences so it is reasonable to believe that all organ systems act in coordination. What has been reviewed so far will be used to exemplify some global properties of each system and argue to what extent the organismal level is to be considered the domain where behavior occurs in a more evident way. The integrative systems (e.g. the nervous and endocrine system) and some other will help us illustrate our view. While some of them overlap, we will focus on the most important elements for pedagogic purposes. Breathing in Pulmonary respiration provides oxygen to the tissues of the organism and remove carbon dioxide via exhalation. Inhalation of air rich in oxygen into the body starts in the nasal passage and is completed from the alveoli up to the nostrils and then out of the body. The structure of the respiratory system is suited to its primary function: transportation of gases in and out of the organism (Barret and Ganong, 2019). The basic composition of the lungs can be identified with pulmonary endothelial cells from which the system develops: Level Cellular Tissular Organal Systemic Organismal Physical substrate Pulmonary endothelial cells Endothelium Lungs Respiratory system Organism Form the inner layer of the lung vasculature Provide a barrier between circulating blood and parenchyma but also perform some metabolic tasks Gas exchange to the bloodstream. They later release carbon dioxide back into the atmosphere Coordinates and integrates the upper and lower respiratory tracts for pulmonary ventilation. Breathing in from oxygen resources found in the immediate environment Function 10 December, 2020 It is to notice that in this example only main organs are considered without neglecting others (e.g. the nasal cavity that filters large particles and humidify air). It is neither limited but superposed with the muscular system as it involves the pharyngeal muscles and the diaphragm. Respiration itself is regulated by the nervous system which adjusts the rate of alveolar ventilation and controls the bronchiolar musculature (Hall, 2016, p. 505). The role of the organism is to breath in from an oxygen-enriched atmosphere by staying outdoors, counting on an artificial respirator, or avoiding a vacuum in space because there is nothing there to inhale. Technically only the lungs respirate or ventilate albeit they cannot inflate without the diaphragm. It is neither the specific function of the individual to breathe, but rather to provide their organs with the external resources needed for the process to begin with. Mating Level Cellular Physical substrate Sperm cells Function Haploid fusion when fertilizing an egg cell Tissular Organic Systemic Organismal Penis, scrotum and testicles Male reproductive system Organism Production and ejaculation of semen Sexual intercourse Mating with an available partner (Cannot divide) 11 Sexual acts can be enhanced by signals such as erotic stimuli which can be provided by imagination. Yet this second example is a bit deceitful because sexual reproduction usually involves two distinct organisms. The external or internal organs of the male reproductive system are also developed from the intermediate mesoderm while sperm and androgens are produced by the testes which are themselves composed of mesothelial cells. The example illustrates that the process itself requires at least two organisms. Female individuals are not fertilized but their eggs. Conversely, flirting, dating, and mating are directed towards reproduction, but are themselves not performed by the sperm cell but by a human individual (or preferably a duetto). Getting stressed Hormone-behavior relationships are of concern for behavioral endocrinology. In turn, endocrine physiology is concerned with the maintenance of various aspects of homeostasis (Barret and Ganong, 2019). Input (sensory) systems and output (muscular) systems are mediated by an integrator system such as the central nervous system while hormones influence all these systems (Nelson, 2010, p. 97). As molecules, they are chemical messengers released by glands for regulating or controlling bodily functions. December, 2020 Cells are influenced by hormones, so hormones rather belong to a chemical level of organization. The brain regulates the hormonal activity of the body, but hormonal changes may also alter neuronal activity. As the CNS can be considered an internal locus of control, the importance for behavior lies not in the endocrine system as such but in the neuroendocrine supersystem. Level Cellular Tissular Organic Systemic Organismal Physical substrate Endocrine cells Epithelial tissue Hypothalamus and endocrine glands HPA, HPT or HPG axes Organism Function Secrete or inhibit the release of glucagon and insulin. Neuroendocrine cells receive neuronal inputs and release a neurohormone into the bloodstream Arrange secretory cells to form glands Release of epinephrine from the adrenal medulla and norepinephrine from the SNS. CRH or ACTH are later released Neuroendocrine integration of the cascade of hormones for regulating bodily processes Waiting in a traffic gridlock 12 Endocrine cells do not form glands but are rather secreted by them and the hypothalamus connects the endocrine with the nervous system. Regardless, every interacting element leading to the final outcome are represented in their respective levels. Consider a behavior such as waiting in a gridlock when in hurry. As a stressor it can typically elicit an unspecific response. The individual must cope with the demanding environment (i.e. coping strategies in psychological jargon) by relieving stress via exercises, yoga, leisure, relaxing, or changing sleep habits. All these activities are performed within a society holding consumer goods, services, or entertainment industry while processes occurring within the skin involve the coordination of hormone secretion and the neural control of behavior. How does a molecule reaching the brain change gene expression or cellular function is of interest for the physiologists, but behavior analysts deal with signal triggers, responses, and consequences that follow them providing a contextual account of the occurrence of behavior. Behavioral endocrinology best illustrates that reduction strategies should integrate our knowledge of hormones and the influence of behavior-environment relations to hormone concentration in the blood. December, 2020 Moving The vernacular meaning of “behavior” usually confine it to observable motor outputs of an organism. But consider the following case: A tutor is concerned with the temper tantrums of her child so she asks a specialist why her toddler does them. Tantrums seems to fulfill a behavioral function, i.e. receive attention, or getting access to tangibles. But if the specialist offers a neurobiological explanation for something that lies in the environment of the child, he or she will not be providing a proper explanation. It is not because a neuromuscular system that the child tantrums, but through it. For behavior analysts, any kind of engaging with the environment counts as behavior, so there are motor, cognitive, and emotional responses. Each one will presumably diverge in their mechanisms (i.e. motor behavior is related to the neuromuscular system while frontal and cortico-thalamic areas are related to cognition). Motor responses can be considered the most expressive aspects of any activity corresponding to an overt realization, e.g. reflexes, habits, or skills. Level Cellular Tissular Physical substrate Myocites Function Cardiac, skeletal or smooth muscle cells have a basic contractile function. Organic Systemic Organismal Skeletal muscle Musculoskeletal system Organism Through motor neurons and in response to neurotransmitters, the striated muscle contracts in a region. By connecting to the skeleton, it allows movement and the maintenance of posture and position of the whole body. Moving through the environment (Not applicable) Here are not considered cellular origins of bones (osteoblasts, osteoclasts and osteocytes) nor is mentioned that there will be no voluntary movement without the action of the nervous system. But it is implicit that the organism moves in its environment through orientation reflexes or by changing from one place to another. If behavior were reduced to motor responses, behavior analysis would be limited to striated muscle physiology. Approximative or withdrawal behavior within an environment still involves responding to spatial restraints and the skeletal muscle would not suffice to answer why did the chicken cross the road. Thinking Covert responses are usually considered nonbehavioral as if they were of another nature (or even nonphysical). A “computational level” is called upon for accounting for information-processing tasks and behavioral goals in terms of input/output functions as a fancy term for referring to what a system does. It is distinguished from a “implementational level” (i.e. physical structures, operations, or mechanisms); and an 13 December, 2020 “algorithmic level” postulated as a middle term for carrying out “representations”. Design principles articulated in this level provide accounts of what phenomena may be produced compared to what does actually perform them (Bechtel and Shagrir, 2015). Neural mechanisms implement psychological functions although “not every implementation will satisfy the conditions for the algorithm (design)” (p. 321). The wellknown Baddeley-Hitch model of memory illustrates the “algorithmic level” enriched with findings of neurosciences. (See Chai et al., 2018) while the computational level can be represented by a recall function R(x,y) without specifying an algorithm of transformation nor its mechanism. For cognitive psychology, the representational level deals with “structures, rules or mental processes” that causally mediate input/output functions. It is therefore “centralist” for being interested in every event that happens inside the [mind of the] organism. A behavioral (nonmediational) “algorithmic” level may be supplemented by a functional diagram or a neuronal network model. It would be neither a “centralist” nor “peripheralist” model but an “interactive” model. If certain processes cannot be topographically identified with the functions of a specific organ, there is only one place for these in the hierarchy of organization, that is, the organismal level. Putting the CNS as the last example is strategic because it is common to reduce behavior (or “mental processes”) to the action of the brain or some part of it. If the rationale of the argument has been followed so far, it will be seen that even covert responses are better explained relating them to the organism. “Cognition” refers to any process by means of which the sensory income is transformed, reduced, elaborated, stored, recovered, or used involving events such as thinking, imagining or planning. “X does y” is quickly identified with the activity of spinal motor neurons that induce voluntary activity or body movements. It is obvious that jumping a fence involves the musculoskeletal system. But it seems less obvious that “x reminds y” is also something x is currently doing or performing. The brain as an information-processing organ describes global properties of a system. Activities in the nervous system are initiated by experiences that excite sensory receptors but also fulfills an effector role by contracting skeletal muscles, smooth muscle in the internal organs or secreting hormones to many parts of the body (Hall, 2016, p. 577). Integrative functions are sometimes posited for accounting for “higher mental activities” as memory or learning. However it is possible to give a behavioral account of cognitive processes such as the provision of previously selected stimuli guiding present responses; or problem solving as supplementing or manipulating stimuli until a contingency (a question) is satisfied. (See Donahoe and Palmer, 2014). They are context-dependent activities and analyzed in terms of contingential relations. As every biological function, they do have a physiological basis. Memory involves the strengthening of synaptic connections and the contribution of second messenger systems on the prefrontal cortex and the lateral and anterior temporal cortex (Barret and Ganong, 2019). 14 December, 2020 But it is a matter of recognizing the synaptic mediators as mechanisms of behavioral responses without neglecting that these are also guided by the environment. Such is the case that destruction of portions of the cerebral cortex does not necessarily prevent a person from having thoughts at all, but it does reduce their depth and the degree of awareness of the surroundings (Hall, 2016, p. 745). Cognitive events are behaviors of the organism and should be interpreted as another form of engaging with its environment. Level Cellular Tissular Organic Systemic Physical substrate Sensory, motor or interneurons and glial cells Nuclei and nerve tracts Cerebrum, brainstem, cerebellum and spinal cord Central nervous system Function Sensory neurons receive and transmit signals. Motor neurons connect to skeletal and smooth muscle. Interneurons connect spinal motor and sensory neurons. Glia provide structural and metabolic support. Association fibers connect a cortical area within the same hemisphere. Commissural fibers connect it with the corresponding area in the opposite hemisphere. Projection tracts connect the cortex with lower parts of the brain and the spinal cord. The brain processes and coordinates sensory information. It is involved in motor control and regulation of autonomic functions. The cord manages messages from the brain to other parts of the body and from sensory receptors to the brain. It also coordinates reflexes alone. Organismal Organism (Covert) Detect energy changes and control movement Providing with substitute stimuli when studying (Overt) Rehearsal of a lecture Mechanisms at different levels are not denied insofar the modus operandi does not preclude other explanatory variables. Furthermore, all (learned) functions of behavior can be interpreted as involving access to tangibles, escape, avoidance, social attention, or sensory (automatic) stimulation (see Iwata and Dozier, 2008). As behavioral processes are related to habituation, sensitization, discrimination, generalization, stimulus equivalence, or substitution, variables controlling these responses are accounted as context-dependent. Organisms within Contexts Behavior is defined as a set of responses under contextual control so it is identified with a unitary organism-environment system. Skinner (1981) stated that behavior evolved by furthering the interchange between the organism and environment as a biological function like others. But under a much wider range of conditions, responding to novel settings may have required a principle of selection related to the individual development. The idea of each organism differentially responding to the same physical event constitutes 15 December, 2020 a tenet for most contextual approaches to behavior. The context of the organism is its “psychological setting” and constitutes the portion of the environment which can be divided in analytical units even when some are not directly manipulated (Fig 3): Environment Context Organism Figure 3. The organism-in-context depiction simplifies the organism-environment system. Not all physical events are functional but only those that reach the organism and control its behavior. The organism interacts in and with a past and current situational context which conforms a “behavioral stream” for each individual. Learning history and the conditions where a response was trained constitute past selections. A rat can be trained in a spatial-learning paradigm to follow a trail when a red light is on. The rat has its own environment and selectively responds to it and although contextual stimulus or cues do not directly control a response, they can alter the outcome if prepotent enough. The context is inclusive as it encompasses all possible variables that control behavior. Zilio (2020) has argued that mechanicism is compatible with contextualism as the search for explanatory mechanisms can be found in both physiological and environmental settings. How selectionism fares with this? Complex behavior is the outcome of variation and selection. What are selected are stimuli-response relations and retained via rehearsal or supplementary cues enabled by neural circuits (e.g. hippocampal synaptic tagging). The selectionist framework is thus a causal model of explanation, while contextualism sets the analytic concepts and rules for predicting and controlling psychological events in a variety of settings although are closely related concepts (Fox, 2006). Organismal factors are often subsumed as contextual variables, i.e., factors that are present when the learning occurs and should be controlled. A similar strategy is found in neuropsychology where the nervous system is seen either as an independent variable for behavioral disorders; or as a dependent variable for exploring the impact of injury, learning, and development of the brain. Roche and Barnes (1997, p. 611) denounce that considering the organism may astray the legitimate goals of the behavior analyst. For example, hands are not intelligently designed for baking but enabled by the joint action of the musculoskeletal system. Yet it is explained by pointing to environmental 16 December, 2020 contingencies. This is the rationale concerning processes such as thinking, reminding, or imagining which are covert responses although may be enabled by the prefrontal cortex to avoid teleological implications. One can even put the organism in brackets while exploring response-environment interactions but for giving a full account of learning and adaptation we are to supplement our research with biological data fostering both functional and mechanismic explanations. Summary The notion of the “organism as a whole” is a vague theoretical concept as it allegedly refers to emergent functions not properly attributable to any of its parts (Bernat, 2019). True is that the total organism and the organism as a whole are not to be confused as the amputation or loss of an organ may diminish a function or even its quality of life, but the individual remains one. It would be easier to dispense the usage of “organism as a whole” referring to it simply as an organism and considering organismal variables either as bodily conditions or physiological mechanisms albeit a molar perspective is more than evident. It is also meaningless to speak of “half an organism” in the same sense of a brain cross section or a portion of kidney. This is meaningful in an anatomical (or morphological) sense, but not in a functional sense. If the organismal level is linked to the establishment of relations with the environment by integrating and coordinating mechanisms and inner states, the role and nature of the organism discussed here allows for a deepening of our accounts of behavior as a relational property. A science of behavior is after all concerned with the functional relations between an organism and its environment and how neural and hormonal systems mediate these (Moore, 2002). But restricting research to the physiological machinery tends to ignore contextual contingencies whereas behavior of organisms is primarily found in orderly environment-responses relations. The biology which most concerns psychologists (or psychobiologists) is thus organismal biology for explaining how living systems develop and function under an array of circumstances. Behavioral functions are processes under contextual control. This does not imply diminishing the legitimacy of the rest of behavioral sciences, but rather emphasizing that psychological or behavioral analysis seeks to study a specific phenomenon, that is, that of interaction with an environment based on the principles of learning. And although it is just one branch among others, it is not a small thing to say that by adapting to an environment it prepares the conditions for development and natural selection to occur. 17 December, 2020 REFERENCES Baer, D. M. (1976). The organism 98. https://doi.org/10.1159/000271519 as host. Human Development. 19(2): 87– Barrett, K. E., & Ganong, W. F. (2019). Ganong's review of medical physiology. New York, McGraw-Hill Medical. Bechtel, William & Shagrir, Oron. (2015). 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