HANDBOOK OF PSYCHOBIOLOGY

By A. JAIRA

1 EVOLUTIONARY PSYCHOLOGY 1 2 BEHAVIORAL GENETICS 35 3 COMPARATIVE PSYCHOLOGY OF VISION 47 4 COMPARATIVE PSYCHOLOGY OF AUDITION 71 5 COMPARATIVE PSYCHOLOGY OF MOTOR SYSTEMS 109 6 VISUAL PROCESSING IN THE PRIMATE BRAIN 139 7 AUDITORY PROCESSING IN THE PRIMATE BRAIN 187 8 PROCESSING OF TACTILE INFORMATION IN THE PRIMATE BRAIN 211 CONTENTS CHAPTER 1 EVOLUTIONARY PSYCHOLOGY 1 LEVELS OF EXPLANATION IN EVOLUTIONARY PSYCHOLOGY 2 THE METATHEORY LEVEL OF ANALYSIS 3 METATHEORETICAL ASSUMPTIONS THAT ARE CONSENSUALLY HELD BY EVOLUTIONARY SCIENTISTS 3 NATURAL SELECTION 4 ADAPTATION 4 SEXUAL SELECTION 6 INCLUSIVE FITNESS THEORY 7 SPECIAL METATHEORETICAL ASSUMPTIONS OF EVOLUTIONARY PSYCHOLOGY 7 PSYCHOLOGICAL MECHANISMS AS THE MAIN UNIT OF ANALYSIS 8 DOMAIN SPECI CITY OF PSYCHOLOGICAL MECHANISMS 9 THE ENVIRONMENT OF EVOLUTIONARY ADAPTEDNESS 10 THE MIDDLE LEVEL THEORY LEVEL OF ANALYSIS 11 PARENTAL INVESTMENT THEORY 12 GOOD GENES SEXUAL SELECTION THEORY 14 THE HYPOTHESES LEVEL OF ANALYSIS 17 GOOD GENES SEXUAL SELECTION THEORY HYPOTHESES 17 THE PREDICTION LEVEL OF ANALYSIS 19 GOOD GENES SEXUAL SELECTION THEORY PREDICTIONS 20 THE FUTURE OF EVOLUTIONARY PSYCHOLOGY 22 THE IMPACT OF EVOLUTIONARY PSYCHOLOGY 24 FUTURE DIRECTIONS 26 REFERENCES 28 EVOLUTIONARY PSYCHOLOGY IS THE APPLICATION OF THE PRINCIPLES AND KNOWLEDGE OF EVOLUTIONARY BIOLOGY TO PSYCHOLOGICAL THEORY AND RESEARCH ITS CENTRAL ASSUMPTION IS THAT THE HUMAN BRAIN IS COMPRISED OF A LARGE NUMBER OF SPECIALIZED MECHANISMS THAT WERE SHAPED BY NATURAL SELECTION OVER VAST PERIODS OF TIME TO SOLVE THE RECURRENT INFORMATION PROCESSING PROBLEMS FACED BY OUR ANCESTORS SYMONS 1995 THESE PROB LEMS INCLUDE SUCH THINGS AS CHOOSING WHICH FOODS TO EAT NEGOTIATING SOCIAL HIERARCHIES DIVIDING INVESTMENT AMONG OFFSPRING AND SELECTING MATES THE ELD OF EVOLUTIONARY PSY CHOLOGY FOCUSES ON IDENTIFYING THESE INFORMATION PROCESSING PROBLEMS DEVELOPING MODELS OF THE BRAIN MIND MECHANISMS THAT MAY HAVE EVOLVED TO SOLVE THEM AND TESTING THESE MODELS IN RESEARCH BUSS 1995 TOOBY COSMIDES 1992 THE ELD OF EVOLUTIONARY PSYCHOLOGY HAS EMERGED DRA MATICALLY OVER THE LAST 15 YEARS AS INDICATED BY EXPONENTIAL GROWTH IN THE NUMBER OF EMPIRICAL AND THEORETICAL ARTICLES IN THE AREA TABLE 1 1 THESE ARTICLES EXTEND INTO ALL BRANCHES OF PSYCHOLOGY FROM COGNITIVE PSYCHOLOGY E G COSMIDES 1989 SHEPARD 1992 TO DEVELOPMENTAL PSYCHOLOGY E G ELLIS MCFADYEN KETCHUM DODGE PETTIT BATES 1999 WEISFELD 1999 ABNORMAL PSYCHOLOGY E G MEALEY 1995 PRICE SLOMAN GARDNER GILBERT RHODE 1994 SOCIAL PSYCHOLOGY E G DALY WILSON 1988 SIMPSON KENRICK 1997 PERSONALITY PSYCHOLOGY E G BUSS 1991 SULLOWAY 1996 MOTIVATION EMOTION E G NESSE BERRIDGE 1997 JOHNSTON 1999 AND INDUSTRIAL ORGANIZATIONAL PSYCHOLOGY E G COLARELLI 1998 STUDD 1996 THE RST UNDERGRADUATE TEXTBOOK ON EVOLUTIONARY PSYCHOLOGY WAS PUBLISHED IN 1999 BUSS 1999 AND SINCE THEN AT LEAST THREE OTHER UNDERGRADU ATE TEXTBOOKS HAVE BEEN PUBLISHED IN THE AREA BARRETT DUNBAR LYCETT 2002 CARTWRIGHT 2000 GAULIN MCBURNEY 2000 IN THIS CHAPTER WE PROVIDE AN INTRODUCTION TO THE ELD OF EVOLUTIONARY PSYCHOLOGY WE DESCRIBE THE METHODOLOGY THAT EVOLUTIONARY PSYCHOLOGISTS USE TO EXPLAIN HUMAN COGNITION AND BEHAVIOR THIS DESCRIPTION BEGINS AT THE BROADEST LEVEL WITH A REVIEW OF THE BASIC GUIDING ASSUMPTIONS THAT ARE EM PLOYED BY EVOLUTIONARY PSYCHOLOGISTS WE THEN SHOW HOW EVOLUTIONARY PSYCHOLOGISTS APPLY THESE ASSUMPTIONS TO DE VELOP MORE SPECI C THEORETICAL MODELS THAT ARE TESTED IN RE SEARCH WE USE EXAMPLES OF SEX AND MATING TO DEMONSTRATE HOW EVOLUTIONARY PSYCHOLOGICAL THEORIES ARE DEVELOPED AND TESTED 2 EVOLUTIONARY PSYCHOLOGY LEVELS OF EXPLANATION IN EVOLUTIONARY PSYCHOLOGY WHY DO SIBLINGS GHT WITH EACH OTHER FOR PARENTAL ATTEN TION WHY ARE MEN MORE LIKELY THAN WOMEN TO KILL SEXUAL RIVALS WHY ARE WOMEN MOST LIKELY TO HAVE EXTRAMARITAL SEX WHEN THEY ARE OVULATING TO ADDRESS SUCH QUESTIONS EVOLU TIONARY PSYCHOLOGISTS EMPLOY MULTIPLE LEVELS OF EXPLANATION RANGING FROM BROAD METATHEORETICAL ASSUMPTIONS TO MORE SPECI C MIDDLE LEVEL THEORIES TO ACTUAL HYPOTHESES AND PREDICTIONS THAT ARE TESTED IN RESEARCH BUSS 1995 KETELAAR ELLIS 2000 THESE LEVELS OF EXPLANATION ARE ORDERED IN A HIERARCHY SEE FIGURE 1 1 AND CONSTITUTE THE METHODOLOGY THAT EVOLUTIONARY PSYCHOLOGISTS USE TO ADDRESS QUESTIONS ABOUT HUMAN NATURE AT THE TOP OF THE HIERARCHY ARE THE BASIC METATHEORETICAL ASSUMPTIONS OF MODERN EVOLUTIONARY THEORY THIS SET OF GUID ING ASSUMPTIONS WHICH TOGETHER ARE REFERRED TO AS EVOLUTION ARY METATHEORY PROVIDE THE FOUNDATION THAT EVOLUTIONARY SCIENTISTS USE TO BUILD MORE SPECI C THEORETICAL MODELS WE BEGIN BY DESCRIBING A THE PRIMARY SET OF METATHEORETICAL ASSUMPTIONS THAT ARE CONSENSUALLY HELD BY EVOLUTIONARY SCI ENTISTS AND B THE SPECIAL SET OF METATHEORETICAL ASSUMPTIONS THAT DISTINGUISH EVOLUTIONARY PSYCHOLOGY WE USE THE TERM EVOLUTIONARY PSYCHOLOGICAL METATHEORY TO REFER INCLUSIVELY TO THIS PRIMARY AND SPECIAL SET OF ASSUMPTIONS TOGETHER AS SHOWN IN FIGURE 1 1 AT THE NEXT LEVEL DOWN IN THE HIER ARCHY JUST BELOW EVOLUTIONARY PSYCHOLOGICAL METATHEORY ARE MIDDLE LEVEL EVOLUTIONARY THEORIES THESE THEORIES ELABORATE THE BASIC METATHEORETICAL ASSUMPTIONS INTO A PARTICULAR PSY CHOLOGICAL DOMAIN SUCH AS MATING OR COOPERATION IN THIS CHAPTER WE CONSIDER TWO RELATED MIDDLE LEVEL EVOLUTIONARY THEORIES PARENTAL INVESTMENT THEORY AND GOOD GENES SEXUAL TABLE 1 1 GROWTH OF PUBLICATIONS IN THE AREA OF EVOLUTIONARY PSYCHOLOGY AS INDEXED BY THE PSYCINFO DATABASE YEARS OF PUBLICATION NUMBER OF PUBLICATIONSA 1985 1988 4 1989 1992 25 1993 1996 100 1997 2000 231 ANUMBER OF ARTICLES BOOKS AND DISSERTATIONS IN THE PSYCINFO DATABASE THAT INCLUDE EITHER THE PHRASE EVOLUTIONARY PSYCHOLOGY OR EVOLUTIONARY PSYCHO LOGICAL IN THE TITLE IN THE ABSTRACT OR AS A KEYWORD ALL ARTICLES FROM THE JOURNAL OF EVOLUTIONARY PSYCHOLOGY WHICH IS A PSYCHOANALYTIC JOURNAL WERE EXCLUDED EVOLUTIONARY PSYCHOLOGICAL METATHEORY MIDDLE LEVEL THEORIES HYPOTHESES SPECIFIC PREDICTIONS BASIC METATHEORETICAL ASSUMPTIONS OF MODERN EVOLUTIONARY THEORY SPECIAL METATHEORETICAL ASSUMPTIONS OF EVOLUTIONARY PSYCHOLOGY ATTACHMENT THEORY BOWLBY 1969 PARENTAL INVESTMENT THEORY TRIVERS 1972 GOOD GENES SEXUAL SELECTION THEORY INDIVIDUALS WHO MORE FULLY DISPLAY TRAITS INDICATIVE OF HIGH GENETIC QUALITY SHOULD BE HEALTHIER AND IN BETTER CONDITION THAN SHOULD CONSPECIFICS WHO DISPLAY THESE TRAITS LESS FULLY THE FREQUENCY AND TIMING OF FEMALE ORGASM SHOULD VARY IN A MANNER THAT SELECTIVELY FAVORS THE SPERM OF MALES WHO DISPLAY INDICATORS OF HIGH GENETIC QUALITY MALES WHO DISPLAY INDICATORS OF HIGH GENETIC QUALITY SHOULD HAVE MORE SEXUAL PARTNERS AND MORE OFFSPRING MORE SYMMETRICAL INDIVIDUALS SHOULD HAVE BETTER MENTAL AND PHYSICAL HEALTH BETTER IMMUNE SYSTEM FUNCTIONING AND LOWER PARASITE LOADS THAN SHOULD LESS SYMMETRICAL INDIVIDUALS THE TIMING AND FREQUENCY OF ORGASMS BY WOMEN SHOULD BE PATTERNED TO SELECTIVELY RETAIN THE SPERM OF MORE SYMMETRICAL MEN MORE SYMMETRICAL MEN SHOULD HAVE MORE LIFETIME SEXUAL PARTNERS AND MORE EXTRAPAIR SEXUAL PARTNERS THAN SHOULD LESS SYMMETRICAL MEN FIGURE 1 1 THE HIERARCHICAL STRUCTURE OF EVOLUTIONARY PSYCHOLOGICAL EXPLANATIONS ADAPTED FROM BUSS 1995 METATHEORETICAL ASSUMPTIONS THAT ARE CONSENSUALLY HELD BY EVOLUTIONARY SCIENTISTS 3 SELECTION THEORY EACH OF WHICH APPLIES THE ASSUMPTIONS OF EVOLUTIONARY PSYCHOLOGICAL METATHEORY TO THE QUESTION OF REPRODUCTIVE STRATEGIES IN DIFFERENT WAYS THESE MIDDLE LEVEL THEORIES ATTEMPT TO EXPLAIN DIFFERENCES BETWEEN THE SEXES AS WELL AS VARIATION WITHIN EACH SEX IN PHYSICAL AND PSYCHOLOGI CAL ADAPTATIONS FOR MATING AND PARENTING AT THE NEXT LEVEL DOWN ARE THE ACTUAL HYPOTHESES AND PRE DICTIONS THAT ARE DRAWN FROM MIDDLE LEVEL EVOLUTIONARY THEO RIES FIGURE 1 1 A HYPOTHESIS IS A GENERAL STATEMENT ABOUT THE STATE OF THE WORLD THAT ONE WOULD EXPECT TO OBSERVE IF THE THEORY FROM WHICH IT WAS GENERATED WERE IN FACT TRUE PREDIC TIONS ARE EXPLICIT TESTABLE INSTANTIATIONS OF HYPOTHESES WE CONCLUDE THIS CHAPTER WITH AN EVALUATION OF HYPOTHESES AND SPECI C PREDICTIONS ABOUT SEXUAL BEHAVIOR THAT HAVE BEEN DE RIVED FROM GOOD GENES SEXUAL SELECTION THEORY SPECIAL ATTEN TION IS PAID TO COMPARISON OF HUMAN AND NONHUMAN ANIMAL LITERATURES THE METATHEORY LEVEL OF ANALYSIS SCIENTISTS TYPICALLY RELY ON BASIC ALTHOUGH USUALLY IMPLICIT METATHEORETICAL ASSUMPTIONS WHEN THEY CONSTRUCT AND EVALU ATE THEORIES EVOLUTIONARY PSYCHOLOGISTS HAVE OFTEN CALLED ON BEHAVIORAL SCIENTISTS TO MAKE EXPLICIT THEIR BASIC ASSUMPTIONS ABOUT THE ORIGINS AND STRUCTURE OF THE MIND SEE GIGERENZER 1998 METATHEORETICAL ASSUMPTIONS SHAPE HOW SCIENTISTS GENERATE DEVELOP AND TEST MIDDLE LEVEL THEORIES AND THEIR DE RIVATIVE HYPOTHESES AND PREDICTIONS KETELAAR ELLIS 2000 THESE BASIC ASSUMPTIONS ARE OFTEN NOT DIRECTLY TESTED AFTER THEY HAVE BEEN EMPIRICALLY ESTABLISHED INSTEAD THEY ARE USED AS A STARTING POINT FOR FURTHER THEORY AND RESEARCH NEWTON S LAWS OF MOTION FORM THE METATHEORY FOR CLASSICAL MECHANICS THE PRINCIPLES OF GRADUALISM AND PLATE TECTONICS PROVIDE A METATHEORY FOR GEOLOGY AND THE PRINCIPLES OF ADAPTATION THROUGH NATURAL SELECTION PROVIDE A METATHEORY FOR BIOLOGY SEVERAL SCHOLARS E G BJORKLUND 1997 RICHTERS 1997 HAVE ARGUED THAT THE GREATEST IMPEDIMENT TO PSYCHOLOGY S DEVELOP MENT AS A SCIENCE IS THE ABSENCE OF A COHERENT AGREED UPON METATHEORY A METATHEORY OPERATES LIKE A MAP OF A CHALLENGING CON CEPTUAL TERRAIN IT SPECI ES BOTH THE LANDMARKS AND THE BOUND ARIES OF THAT TERRAIN SUGGESTING WHICH FEATURES ARE CONSISTENT AND WHICH ARE INCONSISTENT WITH THE CORE LOGIC OF THE META THEORY IN THIS WAY A METATHEORY PROVIDES A SET OF POWERFUL METHODOLOGICAL HEURISTICS SOME TELL US WHAT PATHS TO AVOID NEGATIVE HEURISTIC AND OTHERS WHAT PATHS TO PURSUE POSITIVE HEURISTIC LAKATOS 1970 P 47 IN THE HANDS OF A SKILLED RE SEARCHER A METATHEORY PROVIDES A GUIDE AND PREVENTS CERTAIN KINDS OF ERRORS RAISES SUSPICIONS OF CERTAIN EXPLANATIONS OR OBSERVATIONS SUGGESTS LINES OF RESEARCH TO BE FOLLOWED AND PROVIDES A SOUND CRITERION FOR RECOGNIZING SIGNI CANT OB SERVATIONS ON NATURAL PHENOMENA LLOYD 1979 P 18 THE ULTIMATE CONTRIBUTION OF A METATHEORY IS THAT IT SYNTHESIZES MIDDLE LEVEL THEORIES ALLOWING THE EMPIRICAL RESULTS OF A VARIETY OF DIFFERENT THEORY DRIVEN RESEARCH PROGRAMS TO BE EXPLICATED WITHIN A BROADER METATHEORETICAL FRAMEWORK THIS FACILITATES SYSTEMATIC CUMULATION OF KNOWLEDGE AND PROGRES SION TOWARD A COHERENT BIG PICTURE SO TO SPEAK OF THE SUBJECT MATTER KETELAAR ELLIS 2000 METATHEORETICALASSUMPTIONS THAT ARE CONSENSUALLY HELD BY EVOLUTIONARY SCIENTISTS WHEN ASKED WHAT HIS STUDY OF THE NATURAL WORLD HAD REVEALED ABOUT THE NATURE OF GOD BIOLOGIST J B S HALDANE IS REPORTED TO HAVE MADE THIS REPLY THAT HE HAS AN INORDINATE FONDNESS FOR BEETLES HALDANE S RETORT REFERS TO THE EXTRAORDINARY DI VERSITY OF BEETLE SPECIES FOUND THROUGHOUT THE WORLD SOME 290 000 SPECIES HAVE SO FAR BEEN DISCOVERED E O WILSON 1992 BEETLES MOREOVER COME IN A BEWILDERING VARIETY OF SHAPES AND SIZES FROM TINY GLITTERING SCARAB BEETLES BARELY VISIBLE TO THE NAKED EYE TO PONDEROUS STAG BEETLES WITH MAS SIVE MANDIBLES HALF THE SIZE OF THEIR BODIES SOME BEETLES MAKE A LIVING FORAGING ON LICHEN AND FUNGI OTHERS SUBSIST ON A DIET OF BEETLES THEMSELVES THE RICHNESS AND DIVERSITY OF BEETLE SPECIES ARE MIRRORED THROUGHOUT THE BIOLOGICAL WORLD BIOLOGISTS ESTIMATE THAT ANYWHERE FROM 10 TO 100 MILLION DIFFERENT SPECIES CURRENTLY INHABIT THE EARTH E O WILSON 1992 EACH ONE IN SOME RESPECT DIFFERENT FROM ALL OTHERS HOW ARE WE TO EXPLAIN THIS EXTRAORDINARY RICHNESS OF LIFE WHY ARE THERE SO MANY SPECIES AND WHY DO THEY HAVE THE PARTICULAR CHARACTERISTICS THAT THEY DO THE GENERAL PRINCIPLES OF GENETICAL EVOLUTION DRAWN FROM MODERN EVOLUTIONARY THEORY AS OUTLINED BY W D HAMILTON 1964 AND INSTANTIATED IN MORE CONTEMPORARY SO CALLED SELF ISH GENE THEORIES OF GENETIC EVOLUTION VIA NATURAL AND SEXUAL SELECTION PROVIDE A SET OF CORE METATHEORETICAL ASSUMPTIONS FOR ANSWERING THESE QUESTIONS INCLUSIVE TNESS THEORY CON CEPTUALIZES GENES OR INDIVIDUALS AS THE UNITS OF SELECTION SEE DAWKINS 1976 HAMILTON 1964 WILLIAMS 1966 IN CON TRAST MULTILEVEL SELECTION THEORY IS BASED ON THE PREMISE THAT NATURAL SELECTION IS A HIERARCHICAL PROCESS THAT CAN OPER ATE AT MANY LEVELS INCLUDING GENES INDIVIDUALS GROUPS WITHIN SPECIES OR EVEN MULTI SPECIES ECOSYSTEMS THUS MUL TILEVEL SELECTION THEORY IS CONCEPTUALIZED AS AN ELABORATION OF INCLUSIVE TNESS THEORY ADDING THE CONCEPT OF GROUP LEVEL ADAPTATION RATHER THAN AN ALTERNATIVE TO IT D S WILSON