Biocultural Orchestration of Developmental Plasticity Across Levels: the Interplay of Biology and Culture in Shaping the Mind and Behavior Across the Life Span

Biocultural Orchestration of Developmental Plasticity Across Levels: The Interplay of Biology and Culture in Shaping the Mind and Behavior Across the Life Span

Shu-Chen Li Max Planck Institute for Human Development

The author reviews reemerging coconstructive conceptions of development and recent empirical findings of developmental plasticity at different levels spanning several fields of developmental and life sciences. A cross-level dynamic biocultural coconstructive framework is endorsed to understand cognitive and behavioral development across the life span. This framework integrates main conceptions of earlier views into a unifying frame, viewing the dynamics of life span development as occurring simultaneously within different time scales (i.e., moment-to-moment microgenesis, life span ontogeny, and human phylogeny) and encompassing multiple levels (i.e., neurobiological, cognitive, behavioral, and sociocultural). Viewed through this metatheoretical framework, new insights of potential interfaces for reciprocal cultural and experiential influences to be integrated with behavioral genetics and cognitive neuroscience research can be more easily prescribed.

Metaphorically speaking, two related pendulums, one swinging concerted biological and cultural influences (hence, biocultural back and forth from nature to nurture and the other from brain to coconstructivism) in tuning cognitive and behavioral development mind, have been running the clocks of developmental and cogni- throughout the life span. tive inquiries for centuries. Instead of polarizing toward either end, Coconstructivism or the related interactionism per se is not new. this review of recent advances in life and developmental sciences Conceptions of environmental, cultural, and behavioral factors approaches these two age-old debates through the lens of a cross- interacting with the biological inheritance of human development level dynamic biocultural coconstructivism. Details of this concep- have long philosophical traditions. For instance, Tetens (1777) tion will become clear as the article unfolds. It suffices here to assigned extraordinary plasticity to human nature, thus stipulating underscore the key notions of the term: The effects of a series of opportunities for environmental, cultural, and individual regulation interconnected feed-downward (culture- and context-driven) and during life span ontogeny. At the evolutionary phylogenetic level, feed-upward (neurobiology-driven) interactive processes and de- St. George Jackson Mivart (1871) suggested that behavioral velopmental plasticity at different levels (hence, cross-level) are changes and adaptation precede and affect natural selection. Mod- continuously accumulated via the individual’s moment-to-moment ern behavioral researchers of development have also been sensitive experiences (hence, dynamic) so that, together, they implement to the interplay between biology and various specific aspects of the developmental contexts. Conceptually, there is the consensus that individual ontogeny is hierarchically organized within an open This research was sponsored by the Max Planck Society. I am particu- developmental system that has multiple levels of contexts. Conse- larly grateful to Paul B. Baltes. The seed for this article was planted during quently, developmental phenomena need to be investigated by the various conversations with him while collaborating on a presentation at jointly considering interactions between endogenous and exoge- the 108th Annual Convention of the American Psychological Association, nous processes at various levels (e.g., Alexander, 1969; Anastasi, August 2000, Washington, DC. Throughout the process of preparing this 1958; Baltes, 1979; Baltes & Singer, 2001; Bronfenbrenner, 1979; article, I benefited from the many articles Paul B. Baltes suggested to me Bronfenbrenner & Ceci, 1994; Gottlieb, 1976, 1998; Magnusson, and from stimulating conversations on various topics—in particular, his advocation of biocultural coconstructive influences on life span develop- 1988, 1996; Schaie, 1965). ment with biology and culture both playing active roles. I also thank Tania However, a general awareness of coconstructive notions as such Singer for fruitful exchanges on possibilities of cultural influences on the among developmental researchers is not enough in and of itself to neuronal bases of social cognition and emotion and feedback on earlier resolve the nature–nurture and mind–brain controversies. In fact, versions of this article. I would also like to thank Susan Bluck, Elizabeth these debates have again intensified after the publication of two Carhart, Michael Carhart, Alexandra M. Freund, Gilbert Gottlieb, Judith working drafts of the sequence of the human genome in February Glu¨ck, Tilmann Habermas, Steffen Hillmert, Laura Martignon, Katherine 2001 (Lander et al., 2001; Venter et al., 2001). Through the Nelson, Justin Powell, Jacqui Smith, Paul Verhaeghen, and Qi Wang for marketplace of ideas, media and press coverage of the progress in their valuable discussions or comments on earlier versions of the article. I genomic research are now swinging the age-old pendulums in the give special thanks to Julia Delius for her helpful assistance in language mind of the general public toward genetic and brain determinism and style editing. Last but not least, special thanks also go to all the researchers whose work is reviewed in this article. (Kay, 2000; Moore, 2002; Pa¨a¨bo, 2001). Never has a time been so Correspondence concerning this article should be addressed to Shu- in need of reifying biocultural coconstructive conceptions than Chen Li, Max Planck Institute for Human Development, Center for Life today’s genome era. Span Psychology, Lentzeallee 94, Berlin D-14195, Germany. E-mail: Integration of experiential and cultural influences into the day- [email protected] to-day research practices of behavioral genetics (e.g., McGuffin,

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Riley, & Plomin, 2001), neuroscience (Shepherd, 1991), and cog- across levels. Third, I review reemerging coconstructive concep- nitive neuroscience (e.g., Gazzaniga, 2000) requires that interactions and exemplary empirical findings of developmental plasticity tive processes and mechanisms implementing biocultural cocon- at various levels that extend beyond the period of child develop- struction of brain, mind, and behavior at different levels be further ment, specifically highlighting interactive processes and examples specified. The lack of such specifications, in part, has been one of showing reciprocal cultural influences operating on different time the reasons why the nature–nurture and mind–brain debates have scales corresponding to various levels of developmental plasticity. continued (cf. Ingold, 1998; King, 2000). Thus, a further critical Fourth, to address converging evidence from different levels, I aspect of reifying coconstructive conceptions to help integrate present a summary section of examples of sociocultural influences research from related subfields of developmental and life sciences on, and subsequently expressed through, language development is the need for detailed cross-level frameworks with which inter- and processing. This section is specifically given as a focal illus- active processes and developmental plasticity across different lev- tration of the intimate interconnections between interactive pro- els and time scales can be linked and unified. cesses and developmental plasticity at various levels unfolding across different time scales. Last, new insights into possible inter- Goal and Organization of the Review faces for integrating cultural and experiential influences into the current research agenda and paradigms of behavioral genetics and Taken together, an integral whole of biocultural coconstructive cognitive neuroscience are discussed. influences on life span cognitive and behavioral development most likely are implemented by a series of closely interconnected inter- A Triarchic View of Culture: Resource, Process, and active processes and mechanisms across different levels. Although Developmental Relevancy there have been attempts to study the nature–nurture and mind– brain interactions by different strands of developmental and life A comprehensive treatment of anthropological theories of cul- sciences, studying them separately misses the dynamic gestalt of ture is not within the purview of this article (interested readers the open developmental system cutting across multiple levels. This should see Shweder, 1991; Wertsch, 1985; Wertsch, de Rio, & makes the task of integrating cultural and experiential influences Alvarez, 1995). Here, the focus is on a conception that simulta- into the research practices of behavioral genetics and neuro- neously highlights three distinct but related aspects of culture that sciences extremely difficult, as the interconnections between in- are of direct relevance for considering the dynamics of biocultural teractive processes and developmental plasticity across different coconstructive influences on life span development. Previously, levels and time scales are not in the foreground of many current some scholars have criticized the inadequacy of notions of culture coconstructive conceptions. that view it as acting only to supplement and extend biologically To remedy this situation, the task of this review is thus to piece based capacities. The main concern was that such conceptions together reemerging coconstructive conceptions accompanied by could not address the intertwined relations of culture and biology recent findings of developmental plasticity in related subfields of in the course of human phylogeny and individual ontogeny (e.g., life and developmental sciences by outlining a cross-level meta- Geertz, 1973). Sharing this concern, I define culture in the present theoretical framework to integrate and extend existing concep- context as ongoing collective social processes that generate social, tions. A unique contribution of the current framework is that it psychological, linguistic, symbolic, material, and technological highlights how interactive processes and developmental plasticity resources that influence human development via reciprocal mech- at different levels are closely connected to each other and unfold anisms feeding back into individuals’ moment-to-moment micro- across different time scales. Consequently, together they channel genesis, life span ontogeny involving mind–brain interaction, and reciprocal cultural and experiential influences on behavioral, cog- human phylogeny involving culture–gene coevolution. Culture nitive, and brain development throughout the life span. In princi- defined as such simultaneously encompasses three conjoint facets: ple, biocultural coconstructive conceptions are generalizable to resource, process, and developmental relevancy. various domains of human development. In this article, the sub- stantive focus is on cognitive and behavioral development broadly Culture as Socially Inherited Resources defined, including basic perceptual and cognitive processes, autobiographical memory, language, behavioral adaptation, and learn- Culture has most commonly been conceived as a socially inher- ing. Indeed, physical environment sets boundaries on the human ited combination of human-made material artifacts, knowledge, condition on the evolutionary phylogenetic scale (e.g., Diamond, values, and beliefs accumulated from the past of a given social 1997). However, throughout this review, aside from a few exam- group that functions as a body of resources to coordinate current ples involving other species, the focus is on the socioculturally human life with each other and with the physical environment constructed experiential environment, which plays a more direct (e.g., D’Andrade, 1996; Herskovitz, 1948; Tylor, 1874). Such a role in individual human ontogeny. cumulative process often involves the “ratchet effect” (Tomasello, This article contains five main sections. First, a definition of 1999, p. 5) or “cumulative cultural evolution” (Tomasello, Kruger, culture more apt for examining reciprocal biocultural influences on & Ratner, 1993, p. 495), through which modifications to the development is introduced. Second, a cross-level dynamic biocul- inherited body of cultural resources are being progressively made tural coconstructive metatheoretical framework of life span devel- by individuals of the society over time. Consequently, the inherited opment integrating interactive processes and developmental plas- cultural resources become more complex, encompassing a wider ticity across multiple levels and time scales is proposed. In the range of adaptive functions. For instance, paper, other writing subsequent sections, the metatheory is used as a frame to help materials and tools, books, and variants of printed and electronic piece together recent findings from various subfields of develop- documents are examples of inherited cultural resources accumu- mental and life sciences by interconnecting interactive processes lated through cumulative cultural evolution. Together they support BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 173 intellectual activities in the modern world that comprise both of cultural resources together with the social processes generating conceptual (e.g., ideas and knowledge) and material products that them in individual development. Different cultures (and subcul- have resulted from technological innovations of the past (e.g., the tures) can differ in the level of resources (or opportunities) they inventions of written language, paper, and printing techniques). offer for individual development, in profiles of behavioral expres- If only the resource aspect is emphasized—as is the case in most sions, in the constellations and processes of interactive minds, and generic definitions—then culture is merely the static and passive in their receptivity to modifications by individual behaviors and product of past civilization that is used to support the current way new technological advances (Baltes, 2001). of human life. For a conception of culture to join readily with Taking into account these three facets simultaneously (i.e., coconstructive conceptions, the resource-based view of culture resource, process, and developmental relevancy), culture as de- needs to be extended and combined with a process-oriented notion fined here is not only the passive product of socially inherited and a sense of developmental relevancy. resources of human civilizations such as tools, technology, language, knowledge, art, customs, values, and beliefs that are accu- Culture as an Ongoing Social Process mulated over the past. Rather, it is the “cogenerator” of culture– gene coevolution during human phylogeny in the long run; and Besides the aspect of culture involving socially inherited re- together with behavioral, cognitive, and neurobiological mecha- sources, culture also has a process-oriented dimension involving nisms, it is the active “coproducer” of behavioral, cognitive, and ongoing collective social dynamics (i.e., face-to-face interpersonal neurobiological development during individual life span ontogeny interactions, other social interactions derived from dyadic inter- at present. personal interactions, and shared intentionality) that construct the shared social reality in present societies (Berger & Luckmann, Piecing Together Interactive Processes Across Levels: 1967; Searle, 1995). The processing aspect of culture has been A Cross-Level Dynamic Biocultural Coconstructive particularly emphasized in recent research on cognitive engineer- Framework of Development ing of the workplace and interactive minds (e.g., Baltes & Staudinger, 1996; Hutchins, 1995; Kirsh, 1999). For instance, the With the triarchic definition of culture in place, I now present a routines, habits, and performance criteria of a given workplace are cross-level metatheoretical framework (see Figure 1) highlighting not always static across even relatively short periods (i.e., weeks or concerted biocultural coconstructive influences on life span human months); rather, they are constructed online with the intentions, development that are simultaneously embedded within three time thoughts, and behaviors of members in the group affecting each scales (i.e., phylogenetic, ontogenetic, and microgenetic times) other to shape the collective “working culture.” encompassing multiple levels (i.e., sociocultural, behavioral, cog- Bringing the collective social processing aspect of culture to the nitive, and neurobiological). As a series of interactive processes foreground makes obvious the social interactions taking place in embodied in the framework are explained and substantiated the individual’s proximal developmental context as part of the throughout the remaining sections of this article, it will become processes mediating cultural influences on brain, cognitive, and clear that the proposed framework helps with conceptualizing behavioral development. Furthermore, adding the processing as- intimate interconnections among reemerging coconstructive con- pect also highlights another related feature of culture, namely, ceptions and recent empirical evidence of developmental plasticity cultural plasticity. The very processes generating cultural re- across levels and time scales. Here, a brief overview of key notions sources are themselves pliable and are subject to changes and of the framework is first presented. modifications continually arising from mechanisms of social interaction, social learning, and other large-scale changes in tech- Key Conceptions of the Cross-Level Dynamic Biocultural nology, population, and environmental factors (Baltes & Singer, Coconstructive Framework 2001). Individual ontogeny throughout life is seen as a dynamic pro- Culture as Developmentally Relevant cess that is cumulatively traced out by moment-to-moment experiences and activities taking place at the behavioral, cognitive, and One other somewhat implicit but consistent feature in modern neurobiological levels on the microgenetic time scale. These conceptions of culture is its relevancy for individual development. moment-to-moment microgenetic events are couched within (a) For instance, Williams (1973) pointed out that many of these the proximal developmental context involving culturally embed- notions arise from terms referring to processes that help organisms ded social interactions and situations on the life span ontogenetic grow. Similarly, Cole (1999) noted that throughout history, culture time scale and (b) the distal context of culture–gene coevolution entails a concept of promoting development. occurring on the long-term phylogenetic time scale. In line with the contextual views of culture as an integral part of This framework particularly highlights the interconnections be- the “garden for development” (Cole, 1999, p. 81) or the “devel- tween interactive processes and developmental plasticity at differ- opmental niche” (Super & Harkness, 1986, p. 545, 1997; see also ent levels occurring on different time scales. This is represented Gauvain, 1995, p. 25, 1998), the view of culture in the present schematically in Figure 1 by the dynamic joining of the downward article is one of a medium including both resources and the and upward arrows representing interactive processes occurring on processes generating them, which together edify individual devel- different levels and time scales. Feed-downward influences (i.e., opment to minimize the distance between an individual’s attained cultural influences arising from human phylogeny affecting indi- and potential levels of performance (Vygotsky, 1978). Highlight- vidual moment-to-moment experiences at the behavioral, cogni- ing developmental relevancy as distinct from the resource and tive, and neurobiological levels) are implemented through the process aspects of culture emphasizes the active role and function culture–individual, culture–situation, and situation–individual in- 174 LI

Figure 1. Schematic diagram of the cross-level dynamic biocultural coconstructive framework of development, showing that concerted biocultural influences are implemented through interconnected interactive processes and developmental plasticity across levels and time scales. Dark blue downward arrows denote culture–individual interaction; purple downward arrows denote culture–situation interaction; brown downward arrows denote situation– individual interaction; light blue upward arrows denote individual–situation interaction; pink upward arrows denote situation–culture–gene interaction. The subscripts e, p, and t represent the current epoch, life period, and moment-to-moment microgenetic time, respectively. The notations of these subscripts Ϯ 1...iindicate the sequences before (–) or after (ϩ) the current e, p, or t, where i represents an arbitrary number between 1 and infinity (e.g., p – 1 ϭ the previous life period; p ϩ 1 ϭ the next life period; p ϩ i ϭ a given number of life periods after the current life period). BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 175 teractive processes. Feed-upward influences (i.e., neurobiologi- view of the probabilistic-epigenetic framework. On the other hand, cally implemented individual cognition and behavior contributing although the sociocultural contextual approach focuses on cultural to collective social dynamics and cultural change) are imple- influences on social interactions in the individual’s proximal de- mented through the individual–situation, and situation–culture– velopmental context, interactive processes at the genetic and neu- gene interactive processes. Feed-downward and feed-upward in- ronal levels are not addressed. In addition, both approaches do not fluences are dynamically joined across different time scales, thus explicitly consider the cumulative developmental history as an generating an integral whole of biocultural coconstructive influ- important aspect of the developmental context. Contrary to these ences on human development. approaches is the life span theory of development, which advo- No doubt that the feed-upward genetic and neuronal influences cates the culture–biology interplay throughout the life span but have been and will continue to be the prime research emphases of does not address details of the connections between sociocultural, biogenetics and neurosciences in the new genome era. To balance neuronal, and genetic interactive processes. emphases on the two strains of biocultural influences, a further Similarly, there have been some other recent attempts at the specific task, to paraphrase Gauvain (1995), is to help locate the local specifications of interactive processes, each addressing a feed-downward sociocultural influences in development. Viewed subfacet of the developmental system. For instance, some re- through the present framework, cultural influences on moment-to- searchers have focused on the development of memory and cog- moment microgenesis are implemented through closely linked nition in relation to cultural influences on language development, interactive processes across different levels and time scales. Their because parental speech and culture-specific aspects of language effects are continuously being integrated into the individual’s processing constitute children’s social linguistic environment (e.g., ontogenetic trace and interwoven together with neurobiological Mullen & Yi, 1995; Nelson, 1996; Wang, Leichtman, & Davis, influences to shape the individual’s behavioral, cognitive, and 2000). Others have emphasized the roles of interpersonal relation- brain development across the life span. Therefore, not only do ships in mediating sociocultural influences on child development genetic activities and brain functioning offer the necessary bio- at the behavioral (e.g., Reis, Collins, & Berscheid, 2000) and physical reality for individual cognitive and behavioral operations, neurobiological levels (e.g., Siegel, 1999). Some other researchers when considered within the broader context and time frame of have contended that cumulative developmental history constituted culture and biology coevolution, collective cognition and action by moment-to-moment experiences is an important developmental generated from participatory interactions (Nelson, 1996) among context on its own that leads to specification of functions at the individuals create and define culture. Culture, in turn, orchestrates behavioral (e.g., Thelen & Smith, 1994) and neuronal levels (e.g., individual behavioral, cognitive, and brain development and feeds Johnson, 1999, 2001; Karmiloff-Smith, 1998; Kingsbury & Finlay, back to further cultural change and biological evolution. Further- 2001). Still, there have been others advocating that child cognitive more, albeit the focus here is on processes generating coconstruc- (e.g., Bjorklund & Pellegrini, 2000, 2002; Geary & Bjorklund, tive biocultural influences, it should be underscored that individ- 2000) and brain development (Finlay & Darlington, 1995; Kil- uals need not merely be passive recipients of their biological and lackey, 1990; Kingsbury & Finlay, 2001) should be considered cultural inheritances. Rather, individuals can also be active agents within the larger evolutionary context as subjecting to multiple (Bell, 1968; Lerner & Kauffman, 1985) by participating in making shaping forces and selection pressures. adaptive decisions (Gigerenzer, Todd, & the ABC Research Group, 1999) to regulate the way biocultural influences play out in If research continues to be pursued independently with respect their life histories (Baltes, 1997). to different development contexts at these different levels, the dynamic gestalt of the open developmental system involving a series of closely linked interactions and plasticity across develop- Extending Related Coconstructive Conceptions With a mental contexts at different levels on varying time scales will not Unifying Frame be obtainable. Integration of sociocultural influences on develop- Three existing coconstructive approaches are at the foundation ment into the day-to-day research practices of behavioral genetics of the present framework: (a) the sociocultural contextual ap- and neurosciences would also be difficult to reify. If interconnec- proach viewing child development as simultaneously couched tions between interactive processes and developmental plasticity within multiple time scales and heavily involving culturally em- across levels are not readily in view, new insights for potential bedded social interactions (Cole, 1999; Cole & Cole, 1989; Gau- interfaces and experimental designs testing relations between pro- vain, 1995); (b) the probabilistic-epigenetic framework emphasiz- cesses at different levels are hard to conceive. While being aligned ing bidirectional influences among genes, behavior, and with previous approaches, the present proposal extends beyond environment (Gottlieb, 1991, 1998, 2000); and (c) the life span them by integrating their main conceptions (i.e., different time theory of development highlighting exchanges between cultural frames, multiple levels, and cumulative ontogenetic traces) into a and biological resources throughout life (Baltes, 1987; Baltes & unifying frame that treats life span development as occurring Schaie, 1973; Baltes, Staudinger, & Lindenberger, 1999; Birren, simultaneously within multiple time scales and levels. Such inte- 1964; Nesselroade & Reese, 1973). However, because of the gration brings the interconnections between different levels of complexity of the developmental system involving interactions developmental contexts currently being investigated by various among multiple levels and time scales cutting across disciplinary strands of developmental and life sciences to the foreground. This boundaries, research efforts thus far have been local and divi- proposal thus provides a unifying framework for investigating how sional, missing the dynamic gestalt of such interconnections. For interactive processes and developmental plasticity at different lev- instance, on the one hand, although bidirectional gene– els are related to each other and unfold across different time scales, environment influences are emphasized, complex human social implementing conjoint coconstructive biocultural influences on- situational and cultural linguistic contexts are not within the pur- line throughout life span development. 176 LI

Plasticity at All Levels and Reciprocal Cultural Influences behavior and cognition can only be fully appreciated by incorpo- Operating on Different Time Scales rating an understanding of how human beings, endowed with cultural inheritance (e.g., marital and socioeconomic institutions Biocultural coconstructive influences on development as out- and medical technology), modify significant sources of selection in lined above necessarily imply developmental plasticity at different their environment (e.g., reproductive success, disease survival levels. Reviewing recent progress in various subfields of life and rates, and longevity), thereby codirecting subsequent biological developmental sciences, a clear reemerging zeitgeist of cocon- evolution (e.g., Boyd & Richerson, 1985; Deacon, 1997; Durham, structive and related interactive conceptions that are accompanied 1991; Feldman & Laland, 1996; Laland, Odling-Smee, & Feld- by much recent empirical support for developmental plasticity at different levels1 can be readily gleaned (see summary in the man, 2000; Tomasello, 1999). Along the same lines, Gottlieb Appendix). Two examples of the reemerging coconstructivist zeit- (2002) recently contended specifically that behavioral changes geist are highlighted here. In the area of cognitive sciences broadly incurred during development could instigate genetic changes if defined, coconstructivism has been recently reactivated. For in- transgenerational rearing conditions were relatively stable. stance, Clark (2001) wrote, Evolutionary plasticity. Regarding evolutionary plasticity on the one hand, it has been argued both on empirical and theoretical Recent work in neuroscience, robotics, and psychology...stresses grounds that genetic programs and brain organizations reflect the the unexpected intimacy of brain, body, and world and invites us to sociocultural basis of evolution (e.g., Edelman, 1992). For in- attend to the structure and dynamics of extended adaptive systems.... stance, the dairy farming culture selects for adult lactose tolerance While it needs to be handled with some caution, I believe there is much to be learned from this broader vision. The mind itself, if such and leads to populations in dairy farming societies with a high a vision is correct, is best understood as the activity of an essentially percentage (i.e., over 90%) of lactose absorbers (Aoki, 1986; situated brain: a brain at home in its proper bodily, cultural, and Feldman & Cavalli-Sforza, 1989). Or consider as a different ex- environmental niche [italics added]. ( p. 257) ample, Dunbar’s (1993) proposition that the biological evolution of brain encephalization was, in part, driven by the increase of Advocating the necessity and benefits of integrating cultural and social group size and the emergence of language as a more effi- contextual influences into biological research does not stem only cient means for handling complex social interactions. from the viewpoint of behavioral researchers. In the arena of neurobiology, Jean-Pierre Changeux (1985) saw the possibility of Cultural plasticity. On the other hand, culture itself is also “cultural imprint” and stressed the importance of integrating cul- plastic, subject to changes and modifications by individual behav- tural influences into neuroscience research more than 15 years ago: iors, new technology, or population and environmental factors (Baltes & Singer, 2001). For instance, it has been suggested that Writing leaves an impression on the brain, but where? Our lack of individual learning and experience affect the adoption of cultural knowledge here does not allow us much room for speculation. We traits (Durham, 1991). The invention of writing led to other in- might expect that many areas are involved....Butneurological data ventions such as printing, electronic mailing, and electronically are often hard to interpret; moreover, experimentation is difficult, if not impossible. Nevertheless, the diversity of human culture provides distributed databases. Subsequently, these could affect the roles of fantastically rich material [italics added], and there are a few rare individual minds and external memory devices and the forms and situations where nature carried out its own experiments, even provid- styles of knowledge representation and processing, which together ing controls! (p. 244) constitute new cultural facets of information transaction and transmission (Donald, 1991; Hutchins, 1995; Laland et al., 2000). For Developmental plasticity at the various levels corresponds to instance, the popularity of Internet access over the past decade has interactive processes implementing reciprocal cultural influences already changed the style of academic publishing and many other operating on the time scales of human phylogeny, individual life aspects of daily life, such as shopping and banking activities span ontogeny, and moment-to-moment microgenesis. In what (Eriksen, 2001). In addition to technological development, culture follows, I present conceptions of coconstructivism and develop- is also sensitive to other large-scale environmental and population mental plasticity at different levels with exemplary findings from the macro- (i.e., evolutionary and cultural changes) to the micro- factors. For example, an increase over the 20th century of average levels (i.e., neuronal and genetic activities). Interwoven among life expectancy at birth from 45 to 75 years has resulted in the them are sections highlighting examples of interconnected inter- growing number of aging populations in many developed countries active processes implementing reciprocal cultural influences on (Kannisto, 1994). This could lead to modifications in social and three time scales. Data instantiating culture’s reciprocal long-range economic policies concerning retirement that, in turn, could affect effects on culture–gene coevolution are more of the nature of the retirement culture and roles of seniors in societies. historical and population evidence, whereas data supporting socially mediated effects through the proximal developmental con- 1 text or immediate effects on moment-to-moment microgenetic Outside the immediate realm of developmental research, related interactive conceptions, though not necessarily coconstructive in their empha- experiences are of the empirical nature. ses, are also embedded in current theories taking cross-level integrative approaches to memory (Squire & Kandel, 1999), cognition in general (e.g., Cultural and Evolutionary Plasticity Gazzaniga, 2000), cognitive aging (e.g., Braver et al., 2001; S.-C. Li, 2002; S.-C. Li, Lindenberger, & Sikstro¨m, 2001), and social neuroscience (e.g., At the macrolevel of human phylogeny, there is a growing Cacioppo, Berntson, Sheridan, & McClintock, 2000; Ochsner & Lieber- interest among researchers of human evolution in the interaction man, 2001), as well as in different variants of a contextualized approach to between biological evolution and cultural change. Increasingly, situated and embodied cognition and mental universals (Clark, 1999; researchers have argued that evolutionary approaches to human Pylyshyn, 2000; Shepard, 1995). BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 177

Culture’s Long-Range Effects on Biological Evolution Cultural transmission within a social group may homogenize a population’s behavior. Because culturally transmitted traits can Corresponding to the evolutionary and cultural plasticity dis- spread through the population rapidly compared with genetic vari- cussed above are culture’s long-range reciprocal effects operating ants, culture can generate atypically strong selection (Feldman & on the human phylogenetic scale. On this macroscale, these long- Laland, 1996). If cultural changes are rapid relative to transgen- range effects themselves are composites of various feed-downward erational environmental changes, then the rate of evolutionary and feed-upward interactive processes affecting individual ontog- biological changes may be accelerated. Two illustrative examples eny, social dynamics, and cultural change in the current epoch, of culture’s long-range effects affecting evolutionary biological which together modify the sources of natural selection for biolog- changes expressed at the behavioral and physiological level are ical evolution in the next epoch. given below. For instance, Laland et al. (2000) recently extended conven- Dairy farming and lactose tolerance. For instance, through the tional coevolutionary theory by postulating a set of mediating culture–situation and situation–individual interactions (downward mechanisms called niche construction to relate biological and purple and brown arrows in Figure 1, respectively) dairy farming cultural changes to each other. Central to the concept of niche culture and other sociocultural influences on the proximal devel- construction is the capacity of organisms and individuals to modify opmental context, such as socioeconomic factors, could affect and construct the sources of natural selection in their environment infant-feeding customs and the relative ease of getting dairy prod- by means of (a) learning and experience-dependent processes ucts. Infant-feeding customs, in turn, can affect the amount of during individual ontogeny and (b) processes of cultural change. lactose intake and milk drinking behavior early in and throughout These two broad categories of processes of niche construction can an individual’s life on the microgenetic time scale. Mediated be further dissected as the collective effects of a series of con- through collective social dynamics, these effects could then further nected interactive processes across different levels and time scales affect socioeconomic influences on dairy farming and customs of within the cross-level dynamic biocultural coconstructive frame- dairy product intakes (i.e., individual–situation interaction, light work of development. blue upward arrows in Figure 1). Collective social dynamics, in Processes for learning and experience-dependent changes in- turn, could generate cultural changes and feed back to culture– volve, in part, direct cultural influences on an individual’s gene coevolution in the next epoch (i.e., situation–culture–gene moment-to-moment activities and experiences. Through an indi- interaction, pink upward arrows in Figure 1), selecting for individual’s first-hand personal experiences with the physical or for- viduals with high lactose tolerance. In populations with dairy farming traditions, the percentage of lactose absorbers often ex- mal properties of culture-specific artifacts and symbolic tools ceeds 90, whereas in populations without such a tradition it is evolved over the human phylogenetic time (i.e., direct culture– typically below 20 (Aoki, 1986; Feldman & Cavalli-Sforza, 1989). individual interaction, denoted as the dark blue downward arrows Modern urban living and stress resistance. Consider another in Figure 1), culture exerts direct influences on microgenetic example (Diamond, 1997): The development of agriculture did not moment-to-moment activities at the genetic, neuronal, cognitive, only indirectly lead to changes in individuals’ diets and habitats, it and behavioral levels. The phonological and orthographical prop- also led to changes in societal structures with increasing social erties of a language are examples of formal, rather than social group size (i.e., an example of culture–situation interaction), which interactive, properties of a given language that through culture- then affected the psychological and physiological stresses experi- individual interactions could exert direct influences on the percep- enced by humans in modern societies on the momentary and daily tual and neuronal aspects of language processing. There are also bases (i.e., an example of situation–individual interaction). The culture–situation interactive processes (i.e., culture–situation in- increased stress level in modern urban living, in turn, might teraction, denoted as purple downward arrows in Figure 1) chan- influence biological evolutionary changes in the long run. For neling long-term phylogenetic effects to initially shape culturally instance, the high stress level of modern life could modify the embedded social situational contexts involving social customs, selection pressure to select for individuals with greater stress values, interactions, and concepts about individuals and identity. tolerance. Olson (1999) suggested that on the basis of conditional Subsequently, through situation–individual interactive processes beneficial phenotype, a “less-is-more” mechanism (i.e., having (i.e., situation–individual interaction, denoted as brown downward less phenotypic expression of anxiety under stress means that one arrows in Figure 1) the influences of culturally embedded social is more adaptive to high-stress lifestyles) might be invoked in interactions (e.g., parent–child interaction, schooling, and other biological evolution, in which mutations involving the loss or social interactions) on individual life span development are imple- shedding of sensitive genetic reactions to stress (i.e., the loss-of- mented via moment-to-moment experiences and activities taking gene-function mutation) might occur. place on the microgenetic time scale. Besides the above examples of long-range reciprocal cultural Processes for cultural changes involve individual–situation in- influences being supportive for evolutionary biological changes, teractive processes that cumulatively integrate moment-to-moment there can also be counteractive effects. For instance, late-life experiences at the genetic, neuronal, cognitive, and behavioral illnesses, such as Alzheimer’s disease, with a sharp prevalence rate level into an individual’s ontogenetic history (i.e., individual– increase among 90–100-year-olds, illustrate reciprocal environ- situation interaction, denoted as light blue upward arrows in mental and cultural influences that, in part, counteract evolutionary Figure 1); subsequently, processes of collective social dynamics changes. Given that reproductive-fitness-based evolutionary selec- played out in the individual ontogenetic histories of a society’s tion operates primarily in the first half of life, selection benefits members generate further culture–gene coevolution (i.e., decrease with age. Furthermore, this age-related decrease of evo- situation–culture–gene interaction, denoted as pink upward ar- lutionary selection was further enhanced by the fact that environ- rows in Figure 1). mental and cultural contexts up to the early 1900s allowed a life 178 LI expectancy at birth of only about 45 years, so that only a small five words for individuals in this age range. Further evidence amount of people in the population reached very old age. Besides suggests that marginal cognitive plasticity is still preserved, to other biological processes of aging, genes and cells active in late some extent, in very old age, allowing older adults in the “fourth life are more often dysfunctional than those operating early in life. age” (i.e., aged 80 years and older) to benefit from being informed Given that evolutionary selection cannot operate as frequently about the mnemonic technique (e.g., Singer, Lindenberger, & regarding maladaptive phenotypic characteristics expressed in late Baltes, in press). There is also evidence for adaptive behavioral life as in earlier life periods, dysfunctional late-life genes are not plasticity compensating for the decline of processing resources in likely to be selected against during biological evolution (Finch, old age. For instance, falling has a greater nonadaptive conse- 1990; Hayflick, 1998; Martin, Austad, & Johnson, 1996). quence for older than young adults. Older adults’ more frequent use of walking aids rather than memory aids in a dual task Behavioral and Cognitive Plasticity involving memorization and walking, indicating that they were able to adaptively devote less effort to memorization and more to Regarding the next level of the framework, namely, individual concentration on walking so as to keep themselves from falling ontogeny, various theories in the area of developmental psychol- (K. Z. H. Li, Lindenberger, Freund, & Baltes, 2001). These find- ogy have emphasized the malleability of behavior and cognition by ings together indicate that cognitive and behavioral plasticity are cultural, social, and learning factors (see Collins, Maccoby, Stein- among the hallmarks of individual life span development. berg, Hetherington, & Bornstein, 2000; Lerner, 1998, for reviews). For instance, Vygotsky (1978) had already emphasized the role of Culture’s Midrange Effects on Ontogenetic Behavioral social interactions in proximal developmental contexts such as and Cognitive Development parent–child relation, peer relation, and schooling in promoting an individual’s attained level of development toward a higher level of Corresponding to behavioral and cognitive plasticity are cul- potential. More recently, sociocultural contextual approaches have ture’s midrange effects implemented through culture–situation in- been promoted (e.g., Cole, 1999; Cole & Cole, 1989; Gauvain, teractive processes that affect individuals’ behavior and cognition 1995) that focus more specifically on cultural influences affecting during ontogeny. Cross-cultural differences in parental styles, these social interactions and their subsequent mediated effects on schooling, and other norms of social interaction, such as social- individual development. In a related but different vein, rather than ization goals and self-construal, reveal the cultural influences focusing on cultural influences at a higher level, the relationship accumulated thus far over phylogenetic time on social interactions contextual approach (e.g., Reis et al., 2000) examines fine-grained in the current proximal developmental context of an individual details (e.g., emotional and cognitive aspects) of different types of (i.e., culture–situation interaction). Effects of these culturally em- interpersonal relationships and their impact on child development. bedded social interactions on development are then implemented Still other researchers have focused on the linguistic environment by the behavioral, cognitive, neurobiological, and genetic activities as a main facet of culture-specific social interactions in an indi- on the scale of microgenesis involving moment-to-moment expe- vidual’s proximal developmental context (e.g., Engelkamp, 1978; riences and activities (i.e., situation–individual interaction). In the Mullen, 1994; Nelson, 1996). following sections I provide examples of cultural influences on Cross-cultural studies on parenting style (Bornstein, Tal, & activity goals that subsequently affect infant sleeping behavior and Tamis-LeMonda, 1991) found that differential emphases on inter- of cultural influences on schooling that subsequently affect meta- personal versus object orientation mediated through parent–child cognitive styles. interactions in the Japanese and American cultures, respectively, Intergenerational transmission of activity goals and infant affected the types of languages and games (or playing) at which sleeping behavior. Concrete examples of cultural influences im- the toddlers in these two societies performed well. In the aspect of plemented through culture–situation interactive processes shaping schooling, cultural differences in the emphases of teachers’ in- social interactions in the proximal developmental context involve, structions were shown to affect the acquisition of metacognitive for instance, the collective goals and values shared by members of skills such as memory and computational strategies (Rogoff, a given culture affecting the way individuals conceive their own 1990). It has also been suggested that the differential emphases of life goals. This, in turn, affects the organization of daily planning holistic versus analytical thinking in the East Asian and Western and behavior. For instance, cultural shift caused by industrializa- cultures, respectively, are associated with the individuals’ causal- tion led to a reorganization of cultural practices with respect to attribution style. In this case, analytical thinking style was found to time schedules. Thus, one aspect of becoming modern was behav- be related to a greater propensity in American individuals to ioral change toward a greater sensitivity to planning activities in interpret behavior as the direct result of dispositions corresponding advance (Inkeles & Smith, 1974). Such general cultural shifts in to the apparent nature of the behavior itself. The greater tendency goals and values may subsequently be transmitted into individual of East Asians to include background, contextual, and relational behavior and cognitive development via proximal influences in- information in their causal-attribution processes could, in part, be volving parent–child interaction and other social learning mecha- attributed to their holistic thinking style (e.g., Choi, Nisbett, & nisms (Bronfenbrenner & Ceci, 1994; Cole, 1999; Collins et al., Norenzayan, 1999; Norenzayan & Nisbett, 2000). 2000). As an illustration, consider a study by Super and Harkness In terms of cognitive plasticity in old age, memory training (1997) demonstrating how different societies’ cultural practices of studies (Baltes & Kliegl, 1992) showed that older adults (aged timing and scheduling are transmitted and expressed in young from about 60 to 80 years) still displayed a fair amount of cogni- infants’ sleeping patterns through parent–infant interaction and tive plasticity. They could improve memory recall by up to 10 customs of child care. American urban-dwelling infants whose words by using a culture-based mnemonic technique (i.e., method parents had jobs outside the home and lived by tight working of loci), notwithstanding the usual average memory span of about schedules, developed a marked shift toward the adult day–night BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 179 cycle shortly after birth. By the end of the second week, American dependent neuronal plasticity. After Hebb’s proposal, both the urban babies slept an average of about 8.5 hr between 7 p.m. and selective stabilization theory of neuronal epigenesis (Changeux, 7 a.m. In contrast, the longest period of sleep reported in a sample 1985; Changeux & Danchin, 1976) and the neuronal group selec- of rural Kenyan (Kipsigis) infants was only 3 hr. In comparison tion theory (Edelman, 1987; Edelman & Mountcastle, 1978) fur- with American urban culture, rural Kenyan culture places much ther supported the notion that the details of cortical organization less emphasis on strict scheduling. During the day the Kipsigis are affected by developmental and experiential influences. Since babies napped a lot while strapped to their mothers’ backs, accom- then, many empirical findings have shown that various brain panying them going about their daily activities. During the night regions exhibit experience-dependent structural plasticity (i.e., Kipsigis babies slept with their mothers and were nursed whenever changes in synaptic connectivity and the generation of new neu- they were hungry. Therefore, the Kipsigis babies did not need to rons) and changes in functional cortical circuits (i.e., cortical adapt quickly to a rigid adult day–night cycle. Indeed, the Kipsigis functional plasticity). babies’ longest sleep episode increased little over the first 8 Structural plasticity. It has been demonstrated that the brains months of infancy. of rats reared in environments enabling broader ranges of behav- Schooling and metacognitive styles. Besides parent–child and iors and experiences also show more mature synaptic structure other close interpersonal relationships, other forms of social learn- with more synapses per neuron than rats reared in less stimulating ing, such as schooling, also serve as a channel to implement environments (Black & Greenough, 1998; Greenough & Black, cultural influences on metacognitive styles (Rogoff, 1981, 1990). 1992; Rosenzweig, 1996). In humans, it was recently demonstrated For instance, one study (Carraher, Carraher, & Schliemann, 1985) that the overall cortical gyral patterns, though in part affected by found that for Brazilian child street venders, doing arithmetic had genes (e.g., Pennington et al., 2000), could also be influenced by a socioeconomic context with clear business goals. When the nongenetic experiential factors (Bartley, Jones, & Weinberger, context of mathematic activities and the goal of calculations were 1997). Another example focuses on the amygdala, a brain region meaningful with respect to the sociocultural context, the Brazilian that is very much involved in social–cognitive neurodevelopmen- junior street businessmen’s computations were more successful tal disorders (e.g., autism) and is sometimes referred to as the than when they did the pure arithmetic derivations in classroom innate module for the development of social cognition (e.g., settings common in many other societies. Baron-Cohen, 1995) and emotion (e.g., O¨ hman, 2002). Contrary to In other cross-national studies, Carr, Kurtz, Schneider, Turner, the innate notion, although the amygdala nuclear complex devel- and Borkowski (1989) showed that memory strategy instruction ops relatively early in human gestation (Embryonic Day 30 to 50), varied across different education systems and led to cross-national the separate nuclei do not start to differentiate until after birth, differences in memory performance. Specifically, these authors suggesting plasticity in the amygdala’s responses to experiential found that German and American teachers differed in the type of influences (Kordower, Piecinski, & Rakic, 1992). memory strategy instructions they gave in classrooms. This dif- Functional plasticity. Aside from plasticity in synaptic con- ference was paralleled by the German school children’s higher nectivity, structures, and gyral patterns, neuronal processes also levels of memory recall and memory organizational strategies in exhibit functional plasticity. For instance, the sensorimotor cortex terms of sorting and clustering than their American peers, before in blind individuals develops a larger representation corresponding systematic strategy training was administered to both groups. to the Braille-reading finger and shows cross-modal plasticity in Situation–individual interactive processes channeling the above terms of reorganization in the visual cortex for tactile and auditory intergenerational-, interpersonal-, and schooling-mediated cultural information processing (for reviews see Hamilton & Pascual- influences on individual development are executed in moment-to- Leone, 1998; Kujala, Alho, & Na¨a¨ta¨nen, 2000). Or consider an- moment experiences and activities occurring on the microgenetic other example of a reversed phenomenon: In congenitally deaf time scale involving the behavioral, cognitive, and neurobiological individuals, visual processing dominates parts of the cortex that are levels. Research on mechanisms subserving cultural and social usually involved in auditory function (Neville, 1991; Neville & learning through microprocesses of interpersonal interactions, such Lawson, 1987). as the ability to mentally represent and predict other people’s intentions and behavior (i.e., the ability to have a theory or simu- Cumulative Developmental Influences on Neuronal lation of another’s mind) and associated processes of shared gaze Plasticity Across the Life Span and joint attention, has already begun (e.g., Baron-Cohen, 1995; Carpenter, Nagell, & Tomasello, 1998; Frith & Frith, 1999). I One salient aspect of the evidence on neuronal plasticity shows discuss other interactive processes implementing culture’s imme- that cortical plasticity also reflects the influence of cumulative diate effects on different levels of moment-to-moment experiences developmental history across the life span. A developing brain is after first presenting empirical findings of neuronal and genetic more uniform and lacks the specificity of a developed adult brain. plasticity. Accumulating data suggest that the functional specialization of the neocortex is established through subsequent epigenetic interac- Neuronal Plasticity tions with the immediate experiential environment (Changeux, 1985; see also Johnson, 2001; Kolb & Whishaw, 1998; O’Leary, In the field of neurobiology, since the discovery of long-term 1996, for reviews). Thus, experiential and cultural influences leave potentiation (Bliss & Lømo, 1973)—long-lasting increase in the traces in developing brains, capturing developmental effects accu- synaptic strengthening between simultaneously firing pre- and mulated through the individual’s moment-to-moment activities postsynaptic neurons—which affirmed one aspect of Hebb’s and experiences. For instance, recently it has been demonstrated (1949) principle regarding repeated simultaneous firing, neurosci- that face processing is less localized or specialized (i.e., not as entists have been investigating behavior- and experience- differentiated) in infants than in adults. In infants, face processing 180 LI involves both left and right ventral visual pathways, whereas in related potentials (ERP) studies have demonstrated changes in adults face processing primarily involves the right ventral visual cerebral activity in adults following speech discrimination training pathway. While adult brain activity shows specific sensitivity to or visual word or verbal association learning (e.g., Abdullaev & upright human faces, no such sensitivity for face orientation is Posner, 1997; Kraus et al., 1995; McCandliss, Posner, & Givon, observed in young infants (de Haan, Humphreys, & Johnson, 1997). In terms of language impairment, adult patients who had 2002; de Haan, Pascalis, & Johnson, 2002). recovered from aphasia following strokes showed shifts in lan- Furthermore, there is also increasing evidence for cortical plas- guage lateralization. Specifically, typical Wernicke’s aphasia pa- ticity extending beyond the developing brain to all periods of the tients displayed a shift of function to the right hemisphere that was life span (summarized in Table 1). Many recent data show that the long lasting, whereas patients with Broca’s aphasia displayed a adult brain can also adaptively change its structural and functional transient shift to the right hemisphere that was followed by a return organization in response to accumulated developmental history, to left laterality (Thomas, Altenmu¨ller, Marchmann, Kahrs, & thereby reflecting daily experiences and aging. For instance, Dichgans, 1997). Cortical representational restructuring of senso- Maguire et al. (2000) found that given their extensive navigation rimotor cortex after limb amputation (e.g., Merzenich, Crajski, experience, London taxi drivers’ posterior hippocampi, a region of Jenkins, Recanzone, & Peterson, 1991) and compensatory activa- the brain involved in storing spatial representation of the environ- tion of the right prefrontal cortex to preserve language function ment, were significantly larger in comparison to controls of the after brain damage (e.g., Buckner, Corbetta, Schatz, Raichle, & same age. Furthermore, when comparing among drivers, the num- Petersen, 1996) have also been observed in adults. ber of years spent as a taxi driver correlated positively with As for functional plasticity during aging, a series of recent hippocampal volume. These data indicate that the brain continues neuroimaging data suggests that cortical information processing in to possess functional plasticity in adulthood, allowing the posterior different regions of the brain becomes less differentiated with hippocampus to expand regionally to accommodate elaboration of aging. Compared with young adults, who exhibit more clearly environmental spatial representation in individuals who rely lateralized cortical information processing, people in their 60s and heavily on their navigation skills and who have achieved a high beyond showed bilateralized (bihemispheric) activity during mem- level of navigation expertise in a particular environment. There is ory retrieval (e.g., Cabeza, 2002; Cabeza et al., 1997) and during also evidence with respect to language processing: Recent event- both verbal and spatial working memory processing (e.g., Reuter-

Table 1 Experience and Life-History-Dependent Cortical Plasticity Across the Life Span

Study Finding

Childhood

Neville & Bavelier (1998) Early bilinguals (Ͻ 7 years) showed overlapping cortical activation for native and second language, whereas late bilinguals did not. Elbert et al. (1995) Individuals who learned to play string instruments early showed enlarged cortical representation of the left playing hand. Cheour et al. (1998) Language-specific memory traces developed in infants at 12 months.

Adulthood

Hamilton & Pascual-Leone (1998) Blind individuals developed larger sensorimotor representation corresponding to the Braille-reading finger. There is evidence of recruiting in the visual cortex for tactile information processing. Maguire et al. (2000) The size of posterior hippocampi was enlarged in individuals, such as taxi drivers, who relied heavily on navigation skills in daily activities. Karni et al. (1995) Extensive training of complex finger motor sequences increased the extent of primary motor cortex activation, suggesting slowly evolving, long-term, experience-dependent reorganization of the adult primary motor cortex.

Old age

Cabeza (2002); Reuter-Lorenz et Aging increases bilateral cortical activation in various cognitive al. (2000) processes, indicating reduced hemispheric asymmetry in older adults. Kempermann et al. (1997) Enriched environment induced neurogenesis in the hippocampus of older rats and improved memory performance. H. Nakamura et al. (1999); Saito Enriched environment restores aging-related decrease of et al. (1994) synaptophysin content in older rats by enhanced packing density of synaptic vesicles in synapses.

Note. All data are human data, except two sets of results indicated in the table. BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 181

Lorenz et al., 2000). It has been suggested (Cabeza, 2002; Reuter- Culture’s Immediate Effects on Moment-To-Moment Lorenz, 2002; Reuter-Lorenz et al., 2000) that these data might Experiences indicate that the aging brain could “recruit” cortical areas in both hemispheres to compensate for neurocognitive declines during On the time scale of microgenesis, either through direct culture– aging. individual or mediating culture–situation and situation–individual Regarding structural plasticity in old age, neuroscience’s interactive processes (see Figure 1), culture exerts immediate century-old dogma that there is no addition of new neurons in the effects on moment-to-moment experiences and activities at the adult mammalian brain has recently been revised (see Gross, 2000, behavioral, cognitive, neuronal, and genetic levels. Regarding cul- for a review). There is now evidence showing that increased tural influences implemented through direct culture–individual environmental complexity stimulates the growth of new hip- interactions, the following examples show that individuals’ first- pocampal neurons (i.e., neurogenesis) in the adult brains of various hand personal experiences with the physical or formal properties species, such as birds (Barnea & Nottebohm, 1994), rodents (Kem- of culture-specific artifacts, linguistic, and symbolic tools are permann, Kuhn, & Gage, 1997; Nilsson, Perfilieva, Johannson, possible channels for culture to exert its direct immediate recip- Orwar, & Eriksson, 1999; van Praag, Christie, Sejnowski, & Gage, rocal influences on cognition and behavior. 1999), and humans (Eriksson et al., 1998). Direct influence of culture-specific artifacts on mental calculation. As discussed previously, the metacognitive aspects of math In summary, cultural influences could have effects on the skill learning (i.e., strategy use) involve culture–situation interac- brain’s structural and functional organization through early tive processes mediating cultural influences on educational system experiential tuning of synaptic connections and functional cir- and schooling. The physical properties of culture-specific artifacts cuitry. In addition, though less flexible than in the earlier themselves, however, could have direct effects on moment-to- periods of development, there is still marked cortical plasticity moment experiences affecting online cognitive processing by set- throughout most of the adult life span, both in terms of func- ting specific processing requirements (an example of direct tional plasticity and neurogenesis. This opens up possibilities culture–individual interaction). Consider the example of children for cultural and experiential influences to be intimately inte- skilled at using an abacus (a once common tool of computation grated into an individual’s cumulative developmental history, formerly used daily in East Asian societies) who develop visual- allowing for lifelong adaptations to both ongoing life experi- ized representations of a mental abacus while doing mental calcu- ences couched in the individual’s own respective sociocultural lation. The abacus is a three-dimensional object requiring visual context and life span developmental changes in the efficacy and processing (and also sensorimotor processing at the early acquisi- integrity of the brain itself. tion stage). The visualization of a mental abacus not only enhances the children’s mental calculations but also improves their performance in tasks that require visuospatial processing (Hatano, Mi- Genetic Plasticity yake, & Binks, 1977; Stigler, 1984), such as backward serial recall I now turn to the genetic level of the proposed framework. In the (S.-C. Li & Lewandowsky, 1995). However, at the same time they field of biogenetics research there has been a recent shift from the are also more susceptible to visuospatial distractors (Hatano, traditional view of unidirectional gene3protein information flow Amaiwa, & Shimizu, 1987). (Crick, 1958, 1970) to a probabilistic-epigenetic framework em- Direct influences from formal properties of linguistic or symbolic tools on digit memory and digit and letter processing. phasizing bidirectional interactions among genes, neuronal activ- There is also evidence for linguistically mediated cultural influ- ities, behavior, and environment (Gottlieb, 1991, 1998, 2000). ences on numerical memory span. For instance, the quicker pro- Thus, genetic activity is not viewed as encapsulated and void of nunciation of number words in the Chinese language appears to environmental and behavioral influences; rather, there are recip- contribute to the digit span advantage in Chinese individuals over rocal coactive influences between levels. Environmental and ex- Americans (Geary, Bow-Thomas, Fan, & Siegler, 1993; Stigler, periential influences on genetic activities and expressions have Lee, & Stevenson, 1986). Likewise, more recent experiments show been found in many empirical studies (see summary in the Ap- that besides the ease of pronouncing number words, Chinese pendix; reviews in Gottlieb, 1998). Two examples highlighting preschoolers’ advantage over American preschoolers in the ability developmental plasticity in relation to feed-downward influences to count from 11 to 20 can be related to the more obvious at the genetic level are given here. (10-based) structure of number names in Chinese than in English The acquisition of species-specific songs in songbirds illustrates (Miller, Smith, Zhu, & Zhang, 1995). developmental plasticity implied by feed-downward phylogenetic In addition, other culture-based symbolic tools could also affect influences on moment-to-moment activities at the genetic level. In cognitive processing. For instance, one recent study (Polk & Farah, songbirds, species-specific songs that are present in the environ- 1998) found that even a rather nonsalient aspect of cultural prac- ment stimulate the activation of a set of genes in the forebrain to tice, namely, the composition of postal codes, affected individuals’ further regulate the expression of other genes in the same region letter and digit processing. Canadian postal codes are a mixture of (Mello, Vicario, & Clayton, 1992). Consider also Suomi’s (1999) both digits and letters (e.g., V6K 2E8), and in comparison with finding that maternal style interacted with genetic effects on infant their fellow postal workers who did not sort mail, Canadian mail Rhesus monkeys’ reactions to stressors, which illustrates genetic sorters showed less behavioral evidence for segregated letter and plasticity reflecting the effects of feed-downward influences on digit processing. At the neurobiological level, the study found moment-to-moment activities at the genetic level that are mediated evidence for dissociated digit and letter processing, with the area through parent–child interactions. of left inferior occipitotemporal cortex responding significantly 182 LI more during a letter recognition task than during a digit recogni- the brain are themselves open systems endowed with remarkable tion task. plasticity to be molded by external and experiential influences as An intriguing question is whether there are also traces of cul- individuals behave and function throughout life in their respective tural influences on behavioral and cognitive experiences to be sociocultural environments, which are superimposed on the phys- found at the neurobiological level. The following examples pro- ical environments and embedded in the long-term human phylo- vide initial answers. genetic time scale (cf. Edelman, 2001). Direct cultural influences captured by neuronal plasticity. Be- sides influences channeled through the physical or formal properties of culture-specific artifacts, linguistic, or symbolic tools on the Connecting Interactive Processes Across Levels and Time cognitive level as discussed above, there is evidence showing that Scales: Converging Evidence From Language Research behavioral learning experience with respect to other culture- Although currently available findings regarding cognitive and relevant activities, such as music training, leads to experience- behavioral development as reviewed in the previous section still dependent cortical functional reorganization. Elbert, Pantev, Wien- leave some gaps between levels and time scales, developmental bruch, Rockstroh, and Taub (1995) found that cortical plasticity and interactive processes at the multiple levels may be representations of the fingers of the left playing hands of string more closely related to each other than is readily apparent. Find- players were larger than those of the right hand holding the bow, ings from research on language, unlike those from research on and this was particularly true for individuals who started playing other aspects of cognitive and behavioral development, cover a the instrument early in life. Another more recent neuroimaging wider range of levels and time scales. Converging evidence with study (Ohnishi et al., 2001) compared expert musicians with respect to sociocultural influences on, and expressed through, nonmusicians and found that while listening to J. S. Bach’s Italian language development and processing at most levels and on dif- Concerto, nonmusicians who were not familiar with classical ferent time scales recently has emerged (cf. Kuhl, Tsao, Liu, music showed activity primarily in the secondary auditory associ- Zhang, & de Boer, 2001). In this section, I thus give examples ation area in the right temporal cortex. The musicians, however, involving language to focally illustrate that the various interactive also showed activities in the auditory association area in the left processes and developmental plasticity at different levels as temporal cortex and in the left posterior dorsolateral prefrontal outlined in the cross-level dynamic biocultural coconstructive cortex, the brain regions associated with language processing and framework are tightly related with each other, unfolding across working memory functions, respectively. the time scales of human phylogeny, individual ontogeny, and Parental style- and social interaction–mediated influence cap- microgenesis. tured by genetic plasticity. Feed-downward situation–individual interactive processes (e.g., parent–child interaction and other social situational contexts) implement mediating effects on the indi- Coevolution of Language, Group Size, and Brain: vidual’s moment-to-moment experiences that are, in some cases, Reciprocal Cultural Influence During Human Phylogeny also expressed at the genetic level. Consider again here the example of maternal style interacting with genetic effects on the infant’s Situation–culture–gene and culture–situation interactions oper- reactions to stress. Genetically highly reactive Rhesus monkey ating across the human phylogenetic and individual ontogenetic infants raised by especially nurturant females exhibited much less time scales are both central to the coevolution of language, group deficit in early exploration and less accentuated responses to mild size, and brain. Focusing predominantly on the human phyloge- stressors that were otherwise typical for genetically highly reactive netic scale, Dunbar (1993) showed that group size covaried with infants raised by normal or highly reactive females (Suomi, 1999). relative neocortical volume in nonhuman primates. This led to the Furthermore, evidence has been found of transient, non-learning- proposal that the encephalization of primate brains is not driven by based social psychological stress interacting with mRNA activity the cognitive demands of tool making but by an intricate coevolv- in the human immune system (Glaser et al., 1990), and, for another ing process between the growth in social group size and the example, Malarkey et al. (1996) found that social stressors are development of language as a more efficient method for social related to the concentration of lymphocyte growth hormone in late bonding (Dunbar, 1998). The subsequent effect on the level of adulthood. Specifically, caregivers of spouses with Alzheimer’s individual ontogeny, in turn, was for language to become one of disease showed markedly down-regulated growth hormone gene the socially inherited, species-specific cultural resources that sup- expression. port individuals’ interactions with each other and with the envi- In summary, couched within the context of the cross-level ronment. Indeed, the linguistic relativity theory (Saunders & van dynamic biocultural coconstructive framework, findings of devel- Brakel, 1997; Whorf, 1956) contends that human understanding of opmental plasticity at various levels (i.e., evolutionary, cultural, the world is in part constructed through language. behavioral, cognitive, neuronal, and genetic plasticity) reflect the Brain encephalization is also correlated with the extended juve- effects of coconstructive interactive processes between biology nile period, a phenomenon that possibly also arose from social and culture that dynamically take place across levels and time selection pressures involved in managing complex and dynamic scales. The closely interconnected interactions and plasticity social environments (Bjorklund & Pellegrini, 2000, 2002; Joffe, across levels, thus, form an integral whole of biocultural cocon- 1997). The extended juvenile period in humans, then, allows an structive influences on human development. Together, the existing extended amount of time and opportunities for intergenerational pieces of empirical evidence present a clear warning against the social interactions, operating in conjunction with language, to pure reductionist approach to the genetic and neuronal bases of mediate cultural influences on other aspects of cognitive and mind and behavior, which ignores the influences from cultural, behavioral development during individual ontogeny. To illustrate experiential, and cumulative developmental contexts. Genes and this, I provide a few such examples in the following section. BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 183

Intergenerational Transmission of Self-Construal, Parental manner. As soon as he could speak, he was asked about his former Speech, and Autobiographical Memory: Reciprocal state. But he could not remember any more than we can recall what Cultural Influence During Individual Ontogeny happened to us in the cradle [italics added]. (p. 84) One important aspect of culture–situation interactions on indi- Indeed, recent empirical studies show cross-cultural differences vidual development is the culture-specific language environment, in autobiographical memory, illustrating cultural effects on self- whose effects on an individual’s language and cognitive develop- related memory mediated through the culture-specific social lin- ment are mediated through social interactions. Together with guistic environment. Adults engage children in processes of co- culture-specific characteristics of social interactions, language- constructing memories by explicitly guiding them during specific differences constitute a unique social linguistic environ- conversations, thereby helping them produce verbal accounts of ment as part of the proximal developmental context for individuals their experiences (Eisenberg, 1985; Nelson, 1993). The more in- in different cultures. They affect language and other cognitive terdependent construal of the self in Asian societies seems to relate development during individual ontogeny. Current evidence to a lesser need for elaborated autobiographical narratives, which, shows that cultural differences in terms of how the self is con- in turn, affect early linguistic experiences through parent–child strued (e.g., the Western independent vs. the Asian interdependent interactions. For example, Mullen and Yi (1995) found that the concept of self; Gardner, Gabriel, & Lee, 1999; Markus & frequency with which Korean mother–child dyads engaged in Kitayama, 1991) and language-specific differences in syntax may conversations about past events was only about one third of that of affect the patterns of adult–child interactions, which subsequently U.S. Caucasian mother–child dyads. Wang et al. (2000) compared have implications for language development and autobiographical American and Chinese mother–child dyads and found that the memory. Chinese dyads more often used low-elaborative and interdepen- For instance, Japanese and American children do not differ in dently oriented conversation styles than the American dyads. their language and play skills in general; however, the difference These early linguistic experiences mediated through cultural influ- between the two cultures’ maternal styles of interacting with their ences on self-concept permeate parent–child dynamics and, in infants, either emphasizing more interpersonal dependence (Japa- turn, underlie cross-cultural differences in adult autobiographical nese) or individual independence (American), is related to the memory. The average age of U.S. Caucasian adults’ at their kinds of languages and play skills the infants later develop as earliest childhood memories was about 3.5 years, whereas for the toddlers (e.g., Bornstein et al., 1991). Moreover, another study by Asian adults, it was at about 4 to 4.5 years (Mullen, 1994; Wang, Gopnik, Choi, and Baumberger (1996) shows that language- 2001). specific differences influence parental speech and subsequently Taken together, during individual ontogeny cultural influences affect patterns of semantic and cognitive development. For exam- on an individual’s social linguistic environment that are expressed ple, English has a highly analytic structure, with relative little through cross-cultural differences in intergenerational social dy- reliance on morphological variation. Nouns are generally obliga- namics and language-specific differences affect other aspects of tory in English sentences, and it is rare to construct sentences cognitive and behavioral development. Cross-cultural differences consisting of only an inflected verb without nouns or pronouns. in language syntax influence parents’ style of speech, which later English-speaking parents thus focus heavily on object naming in affects children’s cognitive and semantic abilities (Gopnik et al., conversations with their children. In contrast, the Korean language 1996). Regarding the level of narrative contents, cultural differ- has rich verb morphology, with different verb endings marking ences in how the self is construed affect the content of cocon- important semantic distinctions. Korean parental speech thus often structed narratives between parents and children about the self, consists of highly inflected verbs with few nouns. These authors which subsequently affect the encoding of self-related memory in found that these culture-specific language differences transmitted early childhood (Markus & Kitayama, 1991; Mullen, 1994). The through the parental linguistic environment show subsequent im- fact that this effect later reveals itself as a cross-cultural difference pacts on semantic and cognitive development. Whereas the seman- in the average age of earliest autobiographical memory recalled by tic and cognitive abilities of understanding plans and object cate- individuals in adulthood indeed signifies that cultural and experi- gories developed, on average, at roughly the same age for the ential influences are carried cumulatively throughout the life span. English-speaking children, the Korean-speaking children showed a different pattern. For the Korean-speaking children, means–ends Direct Language-Specific Influences on Moment-To- abilities and verbs that are relevant to the concepts of planning Moment Experiences at Multiple Levels: Reciprocal emerged clearly and consistently before categorization abilities Cultural Influences on the Microgenetic Scale and nouns that are relevant for grouping objects (Gopnik et al., 1996). The above-mentioned socioculturally mediated influences on an In a related vein, there have been historical thoughts and con- individual’s linguistic environment are subsequently implemented temporary studies on the tripartite relation between cultural and by moment-to-moment experiences on the time scale of microgen- experiential influences on language, the concept of self, and auto- esis and may leave traces at the perceptual and neurobiological biographical memory. Consider the following excerpt from a his- levels. Empirical findings corresponding directly to the behavior torical text by Bernard Connor (1697/2001):2 data regarding the style of parental speech, language development, and autobiographical memory are not yet available. Nonetheless, In the forests between Russia and Lithuania a boy of about ten, who lived among the bears, was found in 1694. He gave no sign of reason, walked on hands and feet, had no language, and formed sounds that 2 I thank Michael Carhart at the Max Planck Institute for the History of had nothing in common with human sounds. It took a long time before Science, Berlin, Germany, for pointing out this historical text recounted in he could utter a few words, which he still did in a very barbarous his manuscript of a forthcoming book (Carhart, 2002). 184 LI there are now many data showing the influences of formal prop- dyslexia show better reading performance than English individuals erties of language on moment-to-moment experiences at the per- with dyslexia. ceptual and neuronal levels. Effects of language-specific linguistic environment on language Effects of language-specific linguistic environment on percep- processing captured by neuronal plasticity. Culture-specific lan- tual aspects of language processing. Independent of the social guage differences do not affect language processing at the percep- interaction aspect of the linguistic environment previously dis- tual and cognitive levels only. Recent findings indicate a dynamic cussed, culture-specific differences in formal properties of lan- shift in cortical organization over the course of language acquisi- guage, such as phonology and orthography, also affect language tion (see Neville et al., 1998, for a review). For instance, the time processing. For instance, the attunement model of early experience course of the changes and the degree of experience-dependent on phonological development suggests that culture-specific lan- changes vary with different aspects of language. ERP data indicate that at 20 months, when children are typically speaking in single- guage effects could shape and trim infants’ phonetic discrimina- word utterances, open-class words (i.e., words conveying referen- tion abilities (Aslin, 1981). Young infants (5–17 weeks old) are tial meaning) and closed-class words (i.e., words providing struc- better at discriminating and categorizing phonetic contrasts used in tural and grammatical information) elicit similar patterns of brain different natural languages than adults (e.g., Trehub, 1976). On the activity (Neville & Mills, 1997). At 28 to 30 months of age, when one hand, limiting exposure during early development to the set of children typically begin to speak in short phrases, ERPs to open- phonetic categories of their own native language helps infants and closed-class words reveal different patterns of brain activity, maintain discrimination sensitivity to phonetic contrasts in the and by 3 years of age, when most children speak in sentences and native language, but on the other hand, it leads to loss of sensitivity use closed-class words appropriately like adults, ERPs start to in discriminating sound categories of foreign languages. Thus, display a left hemisphere asymmetry similar to adults (Neville & adults often have difficulty discriminating nonnative phonetic con- Mills, 1997). In a related vein, neuroimaging data indicate strong trast (e.g., Miyawaki et al., 1975; Trehub, 1976). left hemisphere activation for the native language in bilinguals. More recent findings comparing different language groups lend Whereas early bilinguals (i.e., learned their second language be- further support. A study by Kuhl, Williams, Lacerda, Stevens, and fore 7 years of age) showed overlapping areas of cortical activation Lindblom (1992) shows that by 6 months of age, infants start to for the native and second languages (Kim, Relkin, Lee, & Hirsch, discriminate reliably among sound patterns associated with the 1997; Weber-Fox & Neville, 1996), late bilinguals showed inde- language spoken around them but not between the sound patterns pendent or less overlapping activation (see Neville & Bavelier, of unfamiliar languages. Specifically, when American and Swed- 1998; Neville et al., 1998, for reviews). ish infants were tested with vowel prototypes of their native Although language processing depends on a system of neural languages, the infants showed better generalizations to phonetic networks primarily in the left hemisphere, within the common system, there is room for the progressive developmental history of variants around the prototypes contained in their native languages learning and using languages that differ in their orthographical (called the perceptual magnet effect). Another study comparing mapping complexity to leave its trace at the cortical level. Italian two closely related languages also shows that language-specific readers showed greater activation in the left superior temporal phoneme representation is established in infancy and maintained regions, areas associated with phoneme processing, whereas read- throughout adult life (Na¨a¨ta¨nen et al., 1997). Finns and Estonians ers of English showed greater activations—particularly for non- speak closely related languages with very similar vowel structures, words—in the left posterior inferior temporal gyrus and anterior except that the Estonians have one additional vowel /o˜/ which is frontal gyrus, areas associated with word retrieval during both roughly in between the Finnish /o/ (as in sore) and /o¨/ (as in sir). reading and naming tasks (Paulesu et al., 2000). The data seem to Both perceptual and neurophysiological data showed that Finnish suggest that acquiring the rather complex orthographical mapping and Estonian adults judged the vowels common to both languages of the English language impels its readers to invoke additional similarly; however, the response for the /o˜/ category was only neurocognitive mechanisms involving word retrieval from seman- found in the Estonians (Na¨a¨ta¨nen et al., 1997). Furthermore, it was tic memory while reading. found that these language-specific representations emerged before Taken together, examples from language research presented in the age of 12 months (Cheour et al., 1998). this section show that, at a broad level, realizations of the inter- connectedness across levels and time scales unify coconstructive In addition to language-specific phoneme effects, there is also conceptions, interactive processes, and developmental plasticity at evidence of culture-specific language influences operating at the different levels into one general common frame. Viewed through level of orthographical mappings between graphemes and pho- the cross-level dynamic biocultural coconstructive framework, it nemes. In comparison to the Italian language, English orthography becomes explicitly clear that culture–situation, culture–individual, is rather inconsistent, with complicated mappings of letters to situation–individual, individual–situation, and situation–culture– sounds. In the English language there are 1,120 ways of represent- gene interactions jointly played out across the phylogenetic, onto- ing 40 sounds (phonemes), whereas in the Italian language 33 genetic, and microgenetic time scales are all central to the coevo- graphemes are sufficient to represent 25 phonemes. At the lution of language, social dynamics, and brain. While each set of cognitive–perceptual level, this culture-specific language differ- these processes is responsible for separate interactions between ence leads to faster word and nonword reading in Italian university specific levels, as a whole they are closely interconnected; hence, students than in English university students (Paulesu et al., 2000). conceptualizations of various ranges and series of cultural and It is also related to Paulesu et al.’s (2001) finding that the preva- experiential effects on neurobiological processes become more lence rate of dyslexia is about 50% less among Italian than among operant and more easily reified. At a more specific level, in English individuals and that, on average, Italian individuals with keeping with the examples of language research, such realizations BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 185 provide insights for future investigations of social linguistic influ- phenotypic behavioral and cognitive expressions and the proximal ences—such as of sociocultural influences on the development of developmental context (Dickens & Flynn, 2001). self-concept and autobiographical memory or of language-specific Cohort design as a window into history-graded cultural influ- effects mediated through parental speech on semantic develop- ences on genetic heredity. It is conceivable that similar principles ment—at the neurobiological level. could be generalized to examine the interactions between genetic Current research efforts on most other aspects of cognitive and influences and other culture-specific influences in individuals’ behavioral development are still very limited with respect to the proximal developmental contexts. For example, in within-culture levels and time scales they cover. Nonetheless, converging evi- contexts, birth cohort effects are not just influences that need to be dence from studies of language summarized here offers promising controlled for in longitudinal studies. In fact, they may represent prospects for connecting developmental phenomena at various history-graded cultural changes that are primarily driven by tech- levels. Such integration can be achieved by actively incorporating nological developments and changes in population factors, such as considerations of different facets of cultural influences and inter- overall socioeconomic levels. In this vein, studies combining lon- connections between interactive processes across different levels gitudinal cohort-sequential or cross-sequential designs (e.g., unfolding on different time scales into the day-to-day practice of Baltes, 1968; Schaie, 1965) with the emphases of behavioral developmental research. Next, a few new insights into potential genetics research could open up possibilities for exploring ques- interfaces, empirical questions, designs, and methods that would tions on how heredity coefficients might also be influenced by be helpful for integrating cultural and experiential influences with environmental and cultural factors operating within the proximal behavioral genetics and cognitive neuroscience research are developmental contexts of different cohorts. discussed. Recent findings from a population-based Danish twin study (Kohler, Rodgers, & Christensen, 1999) suggest that consider- Insights for Integrating Reciprocal Cultural and ations of demographic regimes shifting the relative magnitudes of Cumulative Developmental Influences Into Behavioral genetic and environmental–cultural contributions to behavior and Genetics and Cognitive Neuroscience Research cognition may not be as irrelevant as they might seem. The authors compared twins born during 1879–1910 and 1953–1964. Al- Overarching dynamic biocultural coconstructive conceptions though genetic influences on fertility existed (estimated to be explicitly detailing the interconnections between interactive pro- about 30%–50% in this sample), the relative patterns were contin- cesses across levels of the kind proposed here are useful in guiding gent on gender and, of particular interest here, on the socioeco- more integrated research efforts across different branches of de- nomic environment experienced by the two cohorts. As genetic velopmental and life sciences. This is a difficult challenge, yet the dispositions may primarily affect fertility behavior and motiva- evidence of reemerging coconstructive views and empirical re- tions for having children, the effect of genetic influence on fertility search at all levels (see Appendix) indicates that, at least, the was accentuated in the later cohort, in which individuals had more possibilities are emerging. Guided by the framework, I discuss in deliberate control over fertility and for which the relatively egal- this last section a few new insights into potential interfaces for itarian socioeconomic opportunities weakened environmental in- reciprocal cultural and cumulative life span developmental influ- fluence across individuals. ences to be integrated into existing behavioral genetics and cog- Life span research as a window into cumulative developmental nitive neuroscience research paradigms. influences on genetic heredity. It is also important to consider the effects of development and aging on the relative balance between Considering Cultural and Cumulative Developmental nature and nurture (Baltes, 1997; Plomin & Crabbe, 2000). For Influences on Genetic Expressions instance, at the descriptive level, heritability of general cognitive ability measured by intelligence tests changes across the life span. Regarding behavioral genetics research, a principle clearly sug- Estimates of heritability are about 20% in infancy, 40% in child- gested by the dynamic biocultural coconstructivism is that genetic hood, 50% in adolescence, and 60% in adulthood and in individ- and environmental–cultural influences interact cumulatively with uals aged 80 years and older (e.g., McClearn et al., 1997; McGue, each other across the life span, jointly affecting behavioral and Bouchard, Iacono, & Lykken, 1993). In addition to cognitive cognitive development. At first glance, this seems a trivial point. abilities, among twins the extent of genetic influences on physical Yet, thus far the traditional “additive model” of behavioral genet- functional abilities in old age was also found to be larger among ics research has treated nature and nurture orthogonally, as if these individuals aged 80 years and older than among individuals two sources of influence added up to 100% of the variance in a aged 75 to 79 years (Christensen et al., 2000). It is interesting that given behavioral or cognitive characteristic. Only recently have concomitant with the increased genetic influences on physical influences on genetic effects by experiential and cultural interac- functional ability in the fourth age (i.e., aged 80 years and older) tions across the life span been emphasized (Baltes, 1997; Dick & is a significant share of environmental influence on older adults’ Rose, 2002; R. J. Rose, 1995) and new research paradigms started health control beliefs, shifting the locus of control from endoge- to emerge. For instance, with respect to cultural and experiential nous self-pertaining factors to exogenous social contextual factors, influences mediated through the proximal developmental context, such as the influence and support from powerful others (Johansson Collins et al. (2000) pointed out that traditional behavioral genetic et al., 2001). Taken together, these findings motivate further ex- research on genetic and parenting effects could be augmented with plorations of the causes and processes, including self-regulatory designs affording either experimental comparisons of differing mechanisms such as motivation and control beliefs, that may parental factors on individuals with varying biological influences underlie developmental and aging effects on the relative contribu- or controls of biological factors. New statistical models have also tions of genetic and environmental influences on cognitive abilities been recently developed to model reciprocal causation between and physical bodily functions in different periods of the life span. 186 LI

Considering Cultural and Cumulative Developmental the interface between the behavioral experimental and cognitive Influences on Cognitive Style and Cortical Functioning neuroscience approaches to aging, expertise, and training research. Take language expertise as an example: comparing the perfor- Regarding cognitive neuroscience research, there are many pos- mance of young and old early and late bilinguals (i.e., individuals sible interfaces for designing research paradigms to more directly who started learning the second language either before and after study the cultural and cumulative life span developmental influ- the age of 7 years, respectively) may help understand the interac- ences on cognitive style and learning- and experience-dependent tions among the age of acquisition, aging effects on expertise in cortical plasticity. In dealing with developmental disorders, language processing, and the related neuronal processes. There Karmiloff-Smith and associates have already argued for the im- have been very few studies comparing young and old bilinguals, portance of considering the possibility of multiple bidirectional and the results show age-related declines in language expertise, mappings from biology to cognition to behavior. Specifically, it particularly in conditions involving switching between two lan- has been suggested that apparent normal behaviors may be guages (Hernandez & Kohnert, 1999). In addition, there seems to achieved by different processes at the cognitive level, which may be a similar decline in both languages, suggesting nonmodular in turn be implemented differently at the neurobiological level. aging effects on first and second language skills (Juncosrabadan, This could be most evident in the case of neurodevelopmental 1994). Given the evidence showing that only early bilinguals disorders because the cognitive–neuronal system has had to work exhibit overlapping cortical activation for first and second lan- around its deficits during development as much as possible (see guages (Neville & Bavelier, 1998), it remains to be seen whether Karmiloff-Smith, 1998, for a comprehensive review). modular (vs. nonmodular) aging effects on language expertise are Neurocognitive aging as a window into cultural and cumulative related to an individual’s age at second-language acquisition and developmental influences on neuronal processes. Neuroimaging the extent of overlapping cortical processing between the data on age-related reduction in lateralization of cognitive infor- languages. mation processing (e.g., Cabeza, 2002; Cabeza et al., 1997; In addition to empirical investigations, neurocomputational Reuter-Lorenz, 2002; Reuter-Lorenz et al., 2000) hint at an anal- models, with their dynamic adaptive properties depending both on ogous picture on cognitive aging. It seems that in response to the input–output mapping contexts and training history, lend aging-related decline in brain efficacy and integrity, “apparent themselves as suitable tools for studying environmental support nominal cognitive tasks” (i.e., tasks with seemingly identical re- (Craik & Anderson, 1999) and experiential and progressive devel- quirements) are implemented differently in the aging brain. It opmental influences on different aspects of cognitive development remains to be seen how much of the cortical functional change and aging. For instance, there are examples of neural network reflects neuronal compensatory functional plasticity in the face of models capturing developmental specifications of language (e.g., losing structural and/or neurochemical integrity; how much re- Elman et al., 1996; Guenther & Gjaja, 1996) and face processing flects the decline in structural, functional, and/or neurochemical (e.g., de Haan, Humphreys, & Johnson, 2002; de Haan, Pascalis, & integrity itself (e.g., Logan, Sanders, Snyder, Morris, & Buckner, Johnson, 2002) in children. Regarding aging, recent neurocompu- 2002); and how much is due to changes in processing strategies at tational theories have been developed to capture the relations the cognitive and behavioral levels. Answers to these questions between aging-related decline in neuromodulation and the dedif- may not be straightforward, however, because these different as- ferentiation (or despecification) of cognitive abilities and other pects are not independent of each other. Neurocomputational mod- aspects of cognitive processing (e.g., S.-C. Li et al., 2001), atten- els could be useful tools for theoretical investigations of these tional control regulation (e.g., Braver et al., 2001), and error intertwined relations (e.g., S.-C. Li et al., 2001; S.-C. Li & Sik- processing (Nieuwenhuis et al., 2002) during aging. stro¨m, 2002). Neurocognitive aging as a window into cultural and cumulative Considering Cultural and Cumulative Developmental developmental influences on cognitive style and expertise acqui- Influences on Neurodevelopmental Disorders sition. In a related vein, recent reports of cultural influences on cognitive styles showing that East Asians process information in a In addition to studies on normative functioning, cultural factors more holistic and contextualized manner whereas Westerners pro- may also be implicated in the behavioral, cognitive, and neuronal cess information in a more analytic and feature-based style (Choi aspects of pathological deficits (Fabrega, 1977; Kennepohl, 1999). et al., 1999; Nisbett, Peng, Choi, & Norenzayan, 2001; Noren- Consider the possibility of examining cultural influences on com- zayan & Nisbett, 2000) suggest that similar cognitive tasks may be ponent processes subserving the development of the ability to processed rather differently by individuals in different cultural predict other individuals’ intentions and behavior (i.e., theory of contexts. Research on how cultural differences might influence the mind), such as attributing desires, engaging in shared attention, relationship between aging and cognition has already begun (Park, and monitoring eye gaze. Although some of these components Nisbett, & Hedden, 1999). Juxtaposing the results from cognitive seem to be related to certain neurobiological substrates and to neuroscience studies on aging and cortical functional lateralization selective impairment in social–cognitive function in individuals with the cognitive–behavioral research on cultural influences on with neurodevelopmental disorders such as autism (see Baron- cognitive style and aging, it seems a possible prospect to integrate Cohen, 1995; Frith & Frith, 1999, for reviews), the question as to designs with both culture-sensitive and culture-invariant cognitive whether the development of these processes could be influenced tasks in the cognitive neuroscience approach to examine how significantly by cultural factors is still open. It is not likely that the environmental and cultural influences interact with the aging pro- functions and developmental sequences of these processes in gen- cesses at the behavioral, cognitive, and neurobiological levels. eral would vary much across different cultures. However, cross- A further possibility for examining cultural influences on cog- cultural variations in whether individual control is considered to be nition and experience-dependent neuronal plasticity seems to lie in internal or external, interpersonal dependent or independent, and in BIOLOGY AND CULTURE IN DEVELOPMENTAL PLASTICITY 187 how necessary it is to explain the internal states of other individ- cultural, experiential, and cumulative developmental contexts) uals are all possible sources of cultural influences on the develop- alone would not help us understand the neurobiological bases of ment of theory of mind (Lillard, 1998). Taking a more domain- mind and behavior. The reason is clear: Genetic activities and relevant (Karmiloff-Smith, 1998) rather than domain-specific neuronal mechanisms themselves possess remarkable plasticity view, it is not implausible that clear cultural differences in cultur- awaiting environment and culture to exert selective developmentally embedded parental styles (e.g., emphasizing interpersonal vs. and experience-based reciprocal influences (cf. Edelman, 1987) on object orientation, interdependent vs. independent self-concept) them and to be the “coauthors” of mind and behavior. People are may interact with the developmental details of these functions, more than mere biological organisms; human mind and behavior which then would need to be considered when applying clinical need to be understood by situating them properly within a brain in assessment instruments and test batteries across countries. a body that lives in an eventful world abounding with objects and I now consider the example of another neurodevelopmental people. Indeed, the brain offers the necessary biophysical reality disorder, Williams syndrome (involving the microdeletion on the for individual cognition and action; it alone, however, is not long arm of chromosome 7 at q11.23), in it has been shown that sufficient to engender the mind or behavior. On the mind–brain there is a within-syndrome double dissociation in terms of uneven continuum, the individual mind is the expression emerging from phenotypic cognitive profiles expressed in infancy and adulthood. the personalized brain (Greenfield, 2000; Llinas & Churchland, Whereas the ability for numerosity judgment is spared in infancy 1996). The very processes for personalizing the biological faculty but impaired in adulthood, language ability is impaired in infancy of the mind take place throughout life span development in envi- but spared in adulthood (Paterson, Brown, Gso¨dl, Johnson, & ronmental and sociocultural contexts, which entail intimate dy- Karmiloff-Smith, 1999). This indicates the importance of includ- namical exchanges between nature and nurture. ing progressive developmental influences into behavioral genetics This review shows that there is accumulating evidence for and cognitive neuroscience research on developmental disorder. developmental plasticity at different levels of analyses that reflect With respect to Williams syndrome, one can also think of analo- and implement, together with closely connected interactive pro- gous implications with respect to including cultural influences into cesses across levels, bidirectional reciprocal biocultural influences the research designs. For instance, the uneven phenotypic cogni- throughout the life span. A cross-level dynamic biocultural cocon- tive profiles of spared verbal fluency and relatively impaired structive framework suggests new insights of possible interfaces spatial-constructive and spatial-perceptual skills in adulthood for the reciprocal cultural influences to be considered and inte- might interact with the differing phonological and orthographic grated into current behavioral genetics and cognitive neuroscience emphases and syntactic and lexical constraints in different lan- research. The task of understanding how individual brains get guages, thereby affecting the hemispheric lateralization of lan- personalized throughout life span development to engender indi- guage processing. To illustrate, the Japanese language has two vidual minds as individuals live together with other sapiens col- systems of signs, the ideograms of Kanji characters and the pho- leagues in a social world abounding with cultural historical heri- netic Kana signs. 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Appendix

Coconstructive Conceptions and Evidence of Developmental Plasticity at Different Levels

Theoretical notions Empirical evidence of plasticity Levels/fields

Coevolution of genes, cognition, and culture Cultural and evolutionary plasticity: receptivity of culture to Evolutionary biology (Dunbar, 1993, 1998; Donald, 1991; Durham, modifications by individual behaviors and new technology and and psychology 1991; Edelman, 1992; Hutchins, 1995) the susceptibility of biological evolution to cognitive and cultural Interactive and coactive evolutionary influences framework between biology, cognition, and Learning and experience influence the adoption of culture traits culture (Durham, 1991). Niche construction Automation and electronically distributed knowledge (Laland et al., 2000) representation and computer-coordinated planning affect the The activities, choices, and metabolic relative roles of individual minds and external memory devices processes through which organisms define, (e.g., Hutchins, 1995). choose, modify, and create their living Invention of writing leads to other inventions (e.g., printing, environment to further modify selection electronic mail; Laland et al., 2000). pressures by feedback to biological Relative neocortical volume in nonhuman primates is related to evolution or cultural changes social group size (Dunbar, 1993, 1998). Development and behavior initiated evolutionary Diary farming selects for lactose tolerance (Aoki, 1986; Feldman change & Cavalli-Sforza, 1989). (Gottlieb, 2002) Incipient specification mediated by developmental–behavioral Behavioral changes incurred during changes in the apple maggot fly (this and other examples are development instigate genetic changes if reviewed in Gottlieb, 2002). transgenerational environmental rearing conditions are relatively stable.

Developmental phenomena need to be studied by Behavioral and cognitive plasticity: socialization and learning- Developmental considering interactional influences at various dependent cognitive and behavioral development across the life psychology levels. The following conceptualizations share the span general notion that individual development is The quality of proximal parental style (e.g., mother–infant hierarchically organized into multiple levels in an interaction) at early ages (2–4 years) affects the amount of open developmental system: problem behavior later in life (Bronfenbrenner & Ceci, 1994; Social-contextual influences Drillien, 1964). (Vygotsky, 1978) The degree of proximal parental monitoring affects school Bioecological model achievement in adolescence (Bronfenbrenner & Ceci, 1994). (Bronfenbrenner & Ceci, 1994) Neighborhood context affects parental style, which subsequently Interactional and holistic framework affects social development in children (Furstenberg, Eccles, (Magnusson, 1988, 1996) Elder, Cook, & Sameroff, 1997). Cultural-social contextual framework Differences in emphasizing interpersonal versus object orientation (Cole, 1999; Cole & Cole, 1989; Gauvain, 1995) in 5-month-old babies in Japan and the United States affect the Biocultural linguistic experiential framework types of languages and playing toddlers are good at (Bornstein (Nelson, 1996; Nelson, Plesa & Henseler, 1998) et al., 1991). Relationship contextual approach Culture affects individuals’ style of causal attribution (Choi et al., (Reis et al., 2000) 1999; Norenzayan & Nisbett, 2000). Interpersonal neurobiology approach Cultural influences on the style of parental speech affect language (Siegel, 1999) development and autobiographical memory (e.g., Gopnik et al., Evolutionary developmental psychology 1996; Mullen, 1994; Wang et al., 2000). (Bjorklund & Pellegrini, 2000, 2002; Geary & At 6 months of age, infants can discriminate sound patterns of Bjorklund, 2000) their own languages (Kuhl et al., 1992). Dynamic system approach Language-specific memory traces develop in infants at the age (Smith & Thelen, 1993; Thelen & Smith, 1994; of 12 months and are maintained throughout adult life (Cheour van Geert, 2000) et al., 1998; Na¨a¨ta¨nen et al., 1997).

Hebbian rule and long-term potentiation (LTP) Neuronal plasticity: behavior- and experience-dependent Cognitive neuroscience (Bliss & Lømo, 1973; Hebb, 1949) synaptogenesis, neurogenesis, and functional plasticity and neurobiology Convergence of firing activities among LTP is important in associative learning and memory. For repeatedly and simultaneously active instance, enduring LTP is observed in hippocampal networks neurons leads to activity-dependent during encoding of associative information (see Squire & strengthening of synaptic connections. Kandel, 1999, for review). Epigenesis by selective stabilization Rats reared in enriched environments showed more mature (Changeux, 1985; Changeux & Danchin, 1976) synaptic structure, more dendritic spines, more synapses per Although genes regulate the division, neuron, better blood supply, more protein content, increased migration, and differentiation of nerve cells, level of neurotransmitters, and growth of new neurons (e.g., epigenetic interactions affect phenotype Black & Greenough, 1998; Greenough & Black, 1992; Gross, variations in fine details of cortical 2000; Rosenzweig, 1996). organization.

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Appendix (continued)

Theoretical notions Empirical evidence of plasticity Levels/fields

Neuronal group selection theory Epigenetic interactions affected the patterns of axonal branching (Edelman, 1987; Edelman & Mountcastle, 1978) in genetically identical clonal water fleas (Changeux, 1985; Although the anatomical structure of the brain Levinthal, Macagno & Levinthal, 1976). is constrained in part by genetic inheritance, Although genetic factors have substantial influences on human cortical organization of cognitive function cerebral size (e.g., Pennington et al., 2000), the overall cortical arises from parallel reciprocal connections gyral patterns, though also affected by genes, are affected by between neuronal groups that are nongenetic factors (Bartley et al., 1997). established by developmental and Enlarged sensorimotor representation of blind individuals’ Braille- experiential selection taking place reading fingers (see Hamilton & Pascual-Leone, 1998, for a throughout life. review). Experience-dependent synaptogenesis Reorganization of cortical sensory representation in terms of (e.g., Greenough & Black, 1992) compensatory plasticity; for instance, binocularly deprived cats Experience-dependent development of synaptic showed improved auditory localization ability (see Rauschecker, connections 1995, for a review). Cortical functional plasticity Cortical representation of the left fingers of stingplayers was (Hamilton & Pascual-Leone, 1998, for a review) larger than that of the right fingers (Elbert et al., 1995). Experience-dependent and compensatory The size of posterior hippocampi was enlarged in individuals changes in functional circuits (e.g., synaptic relying heavily on their navigation skills in daily activities connectivity, functional localization) in (Maguire et al., 2000). various brain regions Dynamic shift in cerebral organization has been found as infants start to learn their native language (Neville et al., 1998).

Probabilistic epigenesis Genetic plasticity: environment and experience-dependent Behavioral genetics and (Gottlieb, 1991, 1998, 2000) genetic activity and expressions biogenetics The production of a functional organism In songbirds, species’ songs stimulate the activation of a set of requires a system with bidirectional genetic, genes in the forebrain to further regulate the expression of other neural, behavioral, and environmental genes in the same region (Mello et al., 1992). influences. In rats, a behavioral learning task involving the vestibular system affected the base nuclear RNA ratios in the vestibular system (Hyde`n & Egyha`zi, 1962). In rats, visual stimulation affected RNA and protein synthesis (particularly RNA diversity) in the visual cortex (Grouse, Schrier, Letendre, & Nelson, 1980; S. P. R. Rose, 1967). In humans, psychological stress affects mRNA activities regarding the immune system (Glaser et al., 1990). Maternal style affects gene expression in reaction to stressors in Rhesus monkeys (Suomi, 1999). In humans, social stressors affect concentration of lymphocyte growth hormone (L-GH) in late adulthood. Caregivers of spouses with Alzheimer’s disease showed markedly suppressed L-GH concentration (Malarkey et al., 1996).

Received April 19, 2001 Revision received June 21, 2002 Accepted June 24, 2002 Ⅲ