Imagine a damp forest floor with a toad sitting motionless by a log. An insect scuttles from under the log, moving too fast to identify, and immediately the toad lunges towards the insect, flicking its tongue towards it. The toad misses its meal this time and the insect swivels away from the lunge of the toad and runs for cover; it was a cockroach (Fig. 3.1). These two behaviours, prey capture by toads and escape running by cockroaches, are excellent case studies in neuroethology because both show how it is possible to identify the roles played by individual nerve cells in recognising significant stimuli and triggering appropriate behavioural responses. In these cases, the stimuli require immediate action on the part of the animal. A toad will in fact try to catch and eat any small animal that moves along the ground in front of it; it probably does not hunt for cockroaches in particular. Likewise a cockroach will turn and run away from rapidly accelerating air currents, such as those produced by the sudden strike of any predator including a toad. Toads and cockroaches are not specifically adapted to detect each other, but natural selection has shaped the evolution of effective neuronal mechanisms that enable toads to recognise scuttling insects as a potential meal and cockroaches to escape from predatory assaults. In each behaviour, the animal needs to assimilate sensory information rapidly and to organise its motor response appropriately.

Summary

The striatum is a subcortical structure thought to be important for higher motor functions and reward processing. It is part of a larger system called the basal ganglia (BG) and composed of multiple subregions thought to be functionally heterogeneous. This review provides information and evidence for the role of the striatum in implementing the fixed action pattern of the grooming chain in the rat. The support for the involvement of the dorsolateral striatal subregion involved in the production of this movement sequence is described, and the general functional significance of implementation by striatal circuitry is discussed. Implementation is meant to refer to the ability of local processing within striatal circuits to enable motor action plans to be completed without distraction from competing sensory or motor demands. The idea that the striatum is involved in more than motor functions is developed and evidence for detailed processing of reward outcomes is presented. We introduce the possibility that the general nature of striatal function of “implementing” chains of information crosses different functional boundaries between movement and reward information. For movement plans, the implementation includes enabling motor sequences for appropriate output and for reward plans, the implementation includes enabling reward incentive hierarchies for appropriate outcome choices. These types of functions could rely upon cross-talk among striatal subregions and reveal a possible shared integrative function for the different “loops” of the BG circuitry.

Explaining grammar as meaningful

In the previous chapter I presented linguistic studies that concentrated on the integration of syntax and conceptual semantics. Chomsky (1957) proposed several varieties of formal structure descriptions originally based on standard frameworks of traditional grammar. Jackendoff (1983) demonstrated that an appropriate integration of phonological, syntactic and semantic phenomena requires a reorganization, in which each of the different domains determines its own principles of formal structure description. We have seen that this insight led to three stages in which the domain’s structure descriptions were represented independently but systematically integrated by means of interface relations. The previous chapter finished with a radical revision: The foundations of different domains and their integration should no longer be represented as phenomena in the body external world, that is, the world of things. Instead the external world should be pushed into the mind/brain/body that organizes the ranges of internal feelings and externally oriented perception, action and objective thought.

The reader may ask why many modern schools of linguistics had widely accepted logical formats as guidelines for descriptive representations. He should recall that antiquity and the middle ages already understood grammar and logic as related disciplines. All other disciplines had to be understood in domain-appropriate conception frameworks following schematic knowledge of grammar and logic. Accounting for the enormous progress of mathematics and logic linguistics also aimed at correspondingly improved formats and theories that nevertheless should be adapted to the characteristics of natural languages. Without abandoning the formalist techniques, theoretical linguistics should clearly circumscribe and define the basic properties that distinguish ordinary language from other notational systems, say of computer science, the genetic code, the “language” of bees etc. or from other knowledge frames defined for the sciences physics, genetics, formal information theory, computer theory etc.

Summary

We explore the possibility that self-grooming in response to the odors or presence of another animal plays a role in olfactory communication. For some animals, the substances released by self-grooming may make groomers more easily detected, more attractive, and/or less threatening to conspecifics that are in close proximity to them. The fact that animals self-groom at different rates when they encounter different individuals suggests that they can target particular conspecifics for purposes of communicating with them. Given that voles and other animals generally spend more time grooming in response to reproductively active, opposite-sex conspecifics than to reproductively quiescent opposite-sex conspecifics, self-grooming may be involved in attracting potential mates and is associated with the behaviors that surround reproduction. Studies have shown that conditions such as endocrine state, diet, age, and familiarity and relatedness of both the groomer and the scent donor affect the amount of time that individuals self-groom when they are exposed to the odors of opposite-sex conspecifics. Consequently, self-grooming in response to the odors of opposite-sex conspecifics may be akin to scent marking in that animals are transmitting odiferous substances into the environment that honestly signal features of their quality and condition to potential mates and competitors.

Introduction

As many terrestrial animals move about their home ranges they are surrounded by scent marks, some are their own and some are those of conspecifics. Animals investigating these scent marks can often determine many features about the individual that deposited them such as its sex, age, reproductive condition, diet, etc.

Grooming and related behaviors

Behavioral and pharmacological research continues to play a crucial role in modern neuroscience, often spearheading new and innovative techniques and models that further our understanding of the intricate workings of the nervous system. This is particularly evident in the arena of mental health research where, with the help of animal models and novel genetic or pharmacological treatments, new insights and theories are evolving to conceptualize more accurately common brain disorders such as anxiety, depression, obsessive–compulsive disorder (OCD), and schizophrenia.

These advances are allowing for an unprecedented examination of the heritable and environmental factors that contribute to disease pathogenesis. However, although there has been marked progress, the biological substrates of many of these disorders remain unclear. To establish a more concrete understanding of these disorders, a careful dissection of experimental phenotypes must be pursued. In this way, every aspect of behavior is a potentially fruitful source of experimental data that can provide clues to the contributing mechanisms.

One important example of such a behavior is grooming. Grooming is a very highly represented behavior in many animals, comprising a large proportion of their waking time. It serves an incredibly diverse range of purposes in the life of the animal from chemocommunication to basic hygiene. It is a natural behavior, yet it can be induced as part of an experimental procedure and has been shown to be sensitive to stress and bidirectionally sensitive to anxiolytic and anxiogenic drugs in rodents, making it an ideal focal point for high-throughput behavioral studies.

The integrated mind/brain/body: a new version of pushing “the world” into the mind/brain/body of a person

The previous chapter finished by emphasizing the importance of archetypes, mentioning also that Langacker’s dominant archetypes belong to the range of objective events. He acknowledged that archetypes in the range of emotions, feelings and self-experience and of other self-experience would transcend his collection of externally observable and objectively analysable facts. In this limited framework important aspects of internal human experience are excluded.

Jackendoff presents a number of arguments for a fundamental extension in which language, thought, perceived things and events in the world are organized in our mind/brain/bodies (Jackendoff 2002, p. 272–273, 305–306). The available structures of the world do not exist independently of our mind/brain/body’s organization generated in mutual cooperation and communication among social groups of people. This is even true for science; theories and measurement techniques are invented, developed, applied and checked in scientific communities. Though there is continuous search for progress, scientific knowledge also is never completed. I agree with Jackendoff that in view of extending our perspective we must go deeper into psychology and neuropsychology of neural assemblies for storing and processing conceptual structures in terms of neural assemblies. They interact with other organization systems, thus generating the interplay and integration of perception, action, attention, selectivity, emotions, feelings and self-awareness and mentalizing the psychology and functional neuropsychology of others. The complete system is in continuous mental and communicative contact with the normal community.

Summary

Electrical stimulation of the midline cerebellum and striatum elicits grooming in rats. Lesioning methods with either surgery or genetic mutations indicate that these brain regions contribute to grooming behaviors. Grid2Lc mutant mice with selective cerebellar atrophy and Girk2Wv mutants with combined cerebellar and substantia nigra atrophy display different effects on grooming. While Grid2Lc mutants were affected in grooming completion but not serial ordering, the reverse was true in Girk2Wv mutants. Our results implicate cerebello–neocortical pathways in the completion of grooming chains, and a striato–pallido–neocortical pathway in the serial ordering of grooming chains.

Introduction and methodological considerations

The role of the cerebellum and basal ganglia on grooming is of some importance considering that grooming implies movement. It is therefore expected that part of the neural circuitry underlying grooming involves some aspect of motor function. In view of the importance of the cerebellum and basal ganglia in balance and posture (Lalonde and Strazielle 2007a), there is a special challenge in interpreting lesion effects of these brain regions on grooming. This is achievable by measuring serial ordering of grooming sequences. It is well established that rodents groom in a cephalocaudal order, anterior before posterior body parts (Richmond and Sachs 1980; Sachs 1988). Different types of grooming components may also be measured, such as face washing; licking of forelimbs, abdomen, back, and hindlimbs; as well as body shaking and scratching (Vanderwolf et al. 1978). Lesions may selectively affect some grooming components in a fashion inexplainable by motor deficits.

Summary

Abnormal phospholipid metabolism has been implicated in the pathogenesis of schizophrenia, and phospholipase C (PLC) β1 was shown to be reduced in specific brain areas of patients with schizophrenia. However, the causal relationship of the PLCβ1 gene with the behavioral symptoms of schizophrenia remains unclear. Recent studies with the knockout (KO) mice for the PLCβ1 gene have revealed an array of interesting phenotypes, which along with other previous information makes the PLCβ1-KO mouse a good candidate for an animal model for schizophrenia. This also suggests that the PLCβ1-linked signaling pathways may be involved in the neural system whose function is disrupted in the pathogenesis of schizophrenia. In this chapter we will introduce various studies relevant to this issue, highlighting the social withdrawal phenotypes of the mutant, such as the lack of barbering behaviors.

Introduction

An animal model for a disease is expected to display endophenotypes, which are quantifiable phenotypes relevant to symptoms of the disease to be modeled (Braff and Freedman 2002; Gould and Gottesman 2006; van den Buuse et al. 2005). The endophenotypes currently pursued in schizophrenia models are: locomotive hyperactivity, sensorimotor gating deficit, deficits in social interaction, and cognitive deficits (e.g., learning and memory). Genetically modified mice targeted on candidate susceptibility genes have so far been generated as animal models for schizophrenia.

Summary

Grooming is increasingly recognized as a reliable marker of stress-related disturbances in animal models of neuropsychiatric disorders. We previously reported that subchronic exposure to 10 mg/kg of phencyclidine (PCP) for 15 days in rats increased grooming expression under both stressful and appetitive conditions, but impaired grooming syntax only when the behavior was elicited with stressful water sprays directed at the face. For the purpose of this chapter, new indexes from the same rats subjected to the water spray condition were analyzed. Results showed that the PCP group aborted less chains after face washing and spent a lower proportion of time in anterior grooming than control animals. Phencyclidine treatment also increased incorrect chain initiations and enhanced the duration of Phase IV within completed syntactic chains. Finally, PCP-injected rats were less engaged in nongrooming activities, and were more inactive. In a context where grooming was needed rostrally after facial contacts with water sprays, these results indicate that subchronic PCP treatment compromised hygiene efficiency and engendered an unfocused and perseverative grooming, most likely combined with an abnormal stress response. These observations suggest that the two leading approaches in the study of grooming patterning may provide pivotal sets of qualitative observations that help identify hygienic and stress-related irregularities in animal models.

The functional triangle of language, mind and brain

In a classical perspective a language consists of the set of its words and sentences determined by its lexicon and grammar. Words and sentences are realized as sound patterns and are mentally registered when we hear them as sound patterns or when we identify them as letter figures on paper. Today they can also be realized and identified as letter configurations on the computer screen.

But there is more. When recalling something said to us, the memorized words and sentences appear as sound images in our minds together with our mental understanding of the words’ and sentences’ meanings. We may also learn that, while our mind thinks, understands, or speaks, some of the grey cells in our brain are active. In a naïve understanding it may appear to us that pieces of uttered words and meanings are realized and kept in the brain like being printed on a physiological tabula rasa or in a storage space.

Many linguists disagree with the assumption that our mind images everything that is relevant for speaking or understanding. They emphasize that when we speak correct language we have no conscious image of all aspects of meaning and the rules that determine grammatical correctness. Indeed for speaking and understanding normal words and sentences the system of grammatical regularities is somehow operative, but we almost never have conscious mental images of them. Thus we must assume that the rules of grammar and the rules of lexical word relations can at best be represented structurally in the manner of an abstract system description. The mental system seems to be similar to other systems of rules such as the well learned intuitive competence of the rules of chess. When we are fluent players of chess we play without consciously concentrating at any moment on the rules.

Some philosophers’ pan-organic outlook: instead of an introduction

The introduction of the first chapter was a plea for the future development of our functional triangle: linguistics–phenomenological psychology–neurocognition. The following illustrations and introductory explanations of brain elements emphasize the enormous challenge of understanding the functional integration of the three disciplines. Serious difficulties confront the interdisciplinary studies due to the differences of empirical methods and different frameworks for principles of thoughts, theorizing and trying to construct fruitful working models. Since this book is not so much concerned with presenting details of measurements and observations the focus is rather on theorizing and clarifying the disciplines’ principles that underlie the conceptual frames for plausible and justified working models.

The differences of interdisciplinary thought are clear. Linguists work with formal constructions considered as plausible and justified working models structuring grammar and systems of meaning rules that describe semantic and pragmatic structure dependencies of lexical words. Their constructions are arrangements of conceptual terms in static relational patterns. The set of terms can only contribute to principles of organization, that is to the topic of the present chapter, when systems of construction rules are added to the collection of terms, rules whose operations identify, combine or separate letter symbols and formal symbol patterns or arrange them in systematically justified patterns representing relations. In modern linguistics these principles of organization have been adapted from organizations introduced in formal logic and formal meta-mathematics during the first half of the past century.