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Monday, May 4, 2020

Deconstructivism

From Wikipedia, the free encyclopedia
 
Deconstructivism is a movement of postmodern architecture which appeared in the 1980s. It gives
the impression of the fragmentation of the constructed building. It is characterized by an absence of harmony, continuity, or symmetry. Its name comes from the idea of "Deconstruction", a form of semiotic analysis developed by the French philosopher Jacques Derrida. Architects whose work is often described as deconstructionism (though in many cases the architects themselves reject the label) include Peter Eisenman, Frank Gehry, Zaha Hadid, Rem Koolhaas, Daniel Libeskind, Bernard Tschumi, and Coop Himmelb(l)au.

Besides fragmentation, Deconstructivism often manipulates the structure's surface skin and creates by non-rectilinear shapes which appear to distort and dislocate elements of architecture. The finished visual appearance is characterized by unpredictability and controlled chaos.

History, context and influences

Deconstructivism came to public notice with the 1982 Parc de la Villette architectural design competition, in particular the entry from Jacques Derrida and Peter Eisenman and the winning entry by Bernard Tschumi, as well as the Museum of Modern Art’s 1988 Deconstructivist Architecture exhibition in New York, organized by Philip Johnson and Mark Wigley. Tschumi stated that calling the work of these architects a "movement" or a new "style" was out of context and showed a lack of understanding of their ideas, and believed that Deconstructivism was simply a move against the practice of Postmodernism, which he said involved "making Doric temple forms out of plywood".

Other influential exhibitions include the 1989 opening of the Wexner Center for the Arts in Columbus, designed by Peter Eisenman. The New York exhibition has featured works by Frank Gehry, Daniel Libeskind, Rem Koolhaas, Peter Eisenman, Zaha Hadid, Coop Himmelb(l)au, and Bernard Tschumi. Since their exhibitions, some architects associated with Deconstructivism have distanced themselves from it; nonetheless, the term has stuck and has come to embrace a general trend within Contemporary architecture.

Modernism and postmodernism


The term Deconstructivism in contemporary architecture is opposed to the ordered rationality of Modernism and Postmodernism. Though postmodernist and nascent deconstructivist architects both published in the journal Oppositions (published between 1973 and 1984), that journal's contents mark a decisive break between the two movements. Deconstructivism took a confrontational stance to architectural history, wanting to "disassemble" architecture. While postmodernism returned to embrace the historical references that modernism had shunned, possibly ironically, deconstructivism rejected the postmodern acceptance of such references, as well as the idea of ornament as an after-thought or decoration.

In addition to Oppositions, a defining text for both deconstructivism and postmodernism was Robert Venturi's Complexity and Contradiction in Architecture (1966). It argues against the purity, clarity and simplicity of modernism. With its publication, functionalism and rationalism, the two main branches of modernism, were overturned as paradigms. The reading of the postmodernist Venturi was that ornament and historical allusion added a richness to architecture that modernism had foregone. Some Postmodern architects endeavored to reapply ornament even to economical and minimal buildings, described by Venturi as "the decorated shed." Rationalism of design was dismissed but the functionalism of the building was still somewhat intact. This is close to the thesis of Venturi's next major work, that signs and ornament can be applied to a pragmatic architecture, and instill the philosophic complexities of semiology. 
 
The deconstructivist reading of Complexity and Contradiction is quite different. The basic building was the subject of problematics and intricacies in deconstructivism, with no detachment for ornament. Rather than separating ornament and function, like postmodernists such as Venturi, the functional aspects of buildings were called into question. Geometry was to deconstructivists what ornament was to postmodernists, the subject of complication, and this complication of geometry was in turn, applied to the functional, structural, and spatial aspects of deconstructivist buildings. One example of deconstructivist complexity is Frank Gehry's Vitra Design Museum in Weil-am-Rhein, which takes the typical unadorned white cube of modernist art galleries and deconstructs it, using geometries reminiscent of cubism and abstract expressionism. This subverts the functional aspects of modernist simplicity while taking modernism, particularly the international style, of which its white stucco skin is reminiscent, as a starting point. Another example of the deconstructivist reading of Complexity and Contradiction is Peter Eisenman's Wexner Center for the Arts. The Wexner Center takes the archetypal form of the castle, which it then imbues with complexity in a series of cuts and fragmentations. A three-dimensional grid, runs somewhat arbitrarily through the building. The grid, as a reference to modernism, of which it is an accoutrement, collides with the medieval antiquity of a castle. Some of the grid's columns intentionally don't reach the ground, hovering over stairways creating a sense of neurotic unease and contradicting the structural purpose of the column. The Wexner Center deconstructs the archetype of the castle and renders its spaces and structure with conflict and difference.

Deconstructivist philosophy

Some Deconstructivist architects were influenced by the French philosopher Jacques Derrida. Eisenman was a friend of Derrida, but even so his approach to architectural design was developed long before he became a Deconstructivist. For him Deconstructivism should be considered an extension of his interest in radical formalism. Some practitioners of deconstructivism were also influenced by the formal experimentation and geometric imbalances of Russian constructivism. There are additional references in deconstructivism to 20th-century movements: the modernism/postmodernism interplay, expressionism, cubism, minimalism and contemporary art. Deconstructivism attempts to move away from the supposedly constricting 'rules' of modernism such as "form follows function," "purity of form," and "truth to materials."

Libeskind's Imperial War Museum North in Trafford, Greater Manchester (2002). An archetype of deconstructivist architecture, it comprises three fragmented, intersecting curved volumes, symbolizing the destruction of war.

The main channel from deconstructivist philosophy to architectural theory was through the philosopher Jacques Derrida's influence with Peter Eisenman. Eisenman drew some philosophical bases from the literary movement Deconstruction, and collaborated directly with Derrida on projects including an entry for the Parc de la Villette competition, documented in Chora l Works. Both Derrida and Eisenman, as well as Daniel Libeskind were concerned with the "metaphysics of presence," and this is the main subject of deconstructivist philosophy in architecture theory. The presupposition is that architecture is a language capable of communicating meaning and of receiving treatments by methods of linguistic philosophy. The dialectic of presence and absence, or solid and void occurs in much of Eisenman's projects, both built and unbuilt. Both Derrida and Eisenman believe that the locus, or place of presence, is architecture, and the same dialectic of presence and absence is found in construction and deconstructivism.

According to Derrida, readings of texts are best carried out when working with classical narrative structures. Any architectural deconstructivism requires the existence of a particular archetypal construction, a strongly-established conventional expectation to play flexibly against. The design of Frank Gehry’s own Santa Monica residence, (from 1978), has been cited as a prototypical deconstructivist building. His starting point was a prototypical suburban house embodied with a typical set of intended social meanings. Gehry altered its massing, spatial envelopes, planes and other expectations in a playful subversion, an act of "de"construction"

In addition to Derrida's concepts of the metaphysics of presence and deconstructivism, his notions of trace and erasure, embodied in his philosophy of writing and arche-writing found their way into deconstructivist memorials. Daniel Libeskind envisioned many of his early projects as a form of writing or discourse on writing and often works with a form of concrete poetry. He made architectural sculptures out of books and often coated the models in texts, openly making his architecture refer to writing. The notions of trace and erasure were taken up by Libeskind in essays and in his project for the Jewish Museum Berlin. The museum is conceived as a trace of the erasure of the Holocaust, intended to make its subject legible and poignant. Memorials such as Maya Lin's Vietnam Veterans Memorial and Peter Eisenman's Memorial to the Murdered Jews of Europe are also said to reflect themes of trace and erasure.

Constructivism and Russian Futurism

Another major current in deconstructivist architecture takes inspiration from the Constructivist and Russian Futurist movements of the early twentieth century, both in their graphics and in their visionary architecture, little of which was actually constructed.

Artists Naum Gabo, El Lissitzky, Kazimir Malevich, and Alexander Rodchenko, have influenced the graphic sense of geometric forms of deconstructivist architects such as Zaha Hadid and Coop Himmelb(l)au. Both Deconstructivism and Constructivism have been concerned with the tectonics of making an abstract assemblage. Both were concerned with the radical simplicity of geometric forms as the primary artistic content, expressed in graphics, sculpture and architecture. The Constructivist tendency toward purism, though, is absent in Deconstructivism: form is often deformed when construction is deconstructed. Also lessened or absent is the advocacy of socialist and collectivist causes.

The primary graphic motifs of constructivism were the rectangular bar and the triangular wedge, others were the more basic geometries of the square and the circle. In his series Prouns, El Lizzitzky assembled collections of geometries at various angles floating free in space. They evoke basic structural units such as bars of steel or sawn lumber loosely attached, piled, or scattered. They were also often drafted and share aspects with technical drawing and engineering drawing. Similar in composition is the deconstructivist series Micromegas by Daniel Libeskind.
The symbolic breakdown of the wall effected by introducing the Constructivist motifs of tilted and crossed bars sets up a subversion of the walls that define the bar itself. ... This apparent chaos actually constructs the walls that define the bar; it is the structure. The internal disorder produces the bar while splitting it even as gashes open up along its length.
— Phillip Johnson and Mark Wigley, Deconstructive Architecture, p. 34

Contemporary art

Two strains of modern art, minimalism and cubism, have had an influence on deconstructivism. Analytical cubism had a sure effect on deconstructivism, as forms and content are dissected and viewed from different perspectives simultaneously. A synchronicity of disjoined space is evident in many of the works of Frank Gehry and Bernard Tschumi. Synthetic cubism, with its application of found object art, is not as great an influence on deconstructivism as Analytical cubism, but is still found in the earlier and more vernacular works of Frank Gehry. Deconstructivism also shares with minimalism a disconnection from cultural references.

With its tendency toward deformation and dislocation, there is also an aspect of expressionism and expressionist architecture associated with deconstructivism. At times deconstructivism mirrors varieties of expressionism, neo-expressionism, and abstract expressionism as well. The angular forms of the Ufa Cinema Center by Coop Himmelb(l)au recall the abstract geometries of the numbered paintings of Franz Kline, in their unadorned masses. The UFA Cinema Center also would make a likely setting for the angular figures depicted in urban German street scenes by Ernst Ludwig Kirchner. The work of Wassily Kandinsky also bears similarities to deconstructivist architecture. His movement into abstract expressionism and away from figurative work, is in the same spirit as the deconstructivist rejection of ornament for geometries. 

Several artists in the 1980s and 1990s contributed work that influenced or took part in deconstructivism. Maya Lin and Rachel Whiteread are two examples. Lin's 1982 project for the Vietnam Veterans Memorial, with its granite slabs severing the ground plane, is one. Its shard-like form and reduction of content to a minimalist text influenced deconstructivism, with its sense of fragmentation and emphasis on reading the monument. Lin also contributed work for Eisenman's Wexner Center. Rachel Whiteread's cast architectural spaces are another instance where contemporary art is confluent with architecture. Ghost (1990), an entire living space cast in plaster, solidifying the void, alludes to Derrida's notion of architectural presence. Gordon Matta-Clark's Building cuts were deconstructed sections of buildings exhibited in art galleries.

1988 MoMA exhibition

Mark Wigley and Philip Johnson curated the 1988 Museum of Modern Art exhibition Deconstructivist architecture, which crystallized the movement, and brought fame and notoriety to its key practitioners. The architects presented at the exhibition were Peter Eisenman, Frank Gehry, Zaha Hadid, Coop Himmelblau, Rem Koolhaas, Daniel Libeskind, and Bernard Tschumi. Mark Wigley wrote the accompanying essay and tried to show a common thread among the various architects whose work was usually more noted for their differences.
The projects in this exhibition mark a different sensibility, one in which the dream of pure form has been disturbed.
It is the ability to disturb our thinking about form that makes these projects deconstructive.
The show examines an episode, a point of intersection between several architects where each constructs an unsettling building by exploiting the hidden potential of modernism.
— Phillip Johnson and Mark Wigley, excerpt from the MoMA Deconstructivist Architecture catalog

Computer-aided design

Computer-aided design is now an essential tool in most aspects of contemporary architecture, but the particular nature of deconstructivism makes the use of computers especially pertinent. Three-dimensional modelling and animation (virtual and physical) assists in the conception of very complicated spaces, while the ability to link computer models to manufacturing jigs (CAM - Computer-aided manufacturing) allows the mass production of subtly different modular elements to be achieved at affordable costs. In retrospect many early deconstructivist works appear to have been conceived with the aid of a computer, but were not; Zaha Hadid's sketches for instance. Also, Gehry is noted for producing many physical models as well as computer models as part of his design process. Though the computer has made the designing of complex shapes much easier, not everything that looks odd is "deconstructivist."

Gallery

Critical responses

Since the publication of Kenneth Frampton's Modern Architecture: A Critical History (first edition 1980) there has been a keen consciousness of the role of criticism within architectural theory. Whilst referencing Derrida as a philosophical influence, deconstructivism can also be seen as having as much a basis in critical theory as the other major offshoot of postmodernism, critical regionalism. The two aspects of critical theory, urgency and analysis, are found in deconstructivism. There is a tendency to re-examine and critique other works or precedents in deconstructivism, and also a tendency to set aesthetic issues in the foreground. An example of this is the Wexner Center. Critical Theory, however, had at its core a critique of capitalism and its excess, and from that respect many of the works of the Deconstructivists would fail in that regard if only they are made for an elite and are, as objects, highly expensive, despite whatever critique they may claim to impart on the conventions of design.

The difference between criticality in deconstructivism and criticality in critical regionalism, is that critical regionalism reduces the overall level of complexity involved and maintains a clearer analysis while attempting to reconcile modernist architecture with local differences. In effect, this leads to a modernist "vernacular." Critical regionalism displays a lack of self-criticism and a utopianism of place. Deconstructivism, meanwhile, maintains a level of self-criticism and a dystopianism of place, as well as external criticism and tends towards maintaining a level of complexity. Some architects identified with the movement, notably Frank Gehry, have actively rejected the classification of their work as deconstructivist.

Critics of deconstructivism see it as a purely formal exercise with little social significance. Kenneth Frampton finds it "elitist and detached". Nikos Salingaros calls deconstructivism a "viral expression" that invades design thinking in order to build destroyed forms; while curiously similar to both Derrida's and Philip Johnson's descriptions, this is meant as a harsh condemnation of the entire movement. Other criticisms are similar to those of deconstructivist philosophy—that since the act of deconstructivism is not an empirical process, it can result in whatever an architect wishes, and it thus suffers from a lack of consistency. Today there is a sense that the philosophical underpinnings of the beginning of the movement have been lost, and all that is left is the aesthetic of deconstructivism. Other criticisms reject the premise that architecture is a language capable of being the subject of linguistic philosophy, or, if it was a language in the past, critics claim it is no longer. Others question the wisdom and impact on future generations of an architecture that rejects the past and presents no clear values as replacements and which often pursues strategies that are intentionally aggressive to human senses.

Animal culture

From Wikipedia, the free encyclopedia

Animal culture involves the current theory of cultural learning in non-human animals, through socially transmitted behaviors. The question as to the existence of culture in non-human societies has been a contentious subject for decades, largely due to the lack of a concise definition for the word "culture". However, many leading scientists agree on seeing culture as a process, rather than an end product. This process, most agree, involves the social transmittance of novel behaviour, both among peers and between generations. Such behaviour can be shared by a group of animals, but not necessarily between separate groups of the same species.

The notion of culture in animals dates back to Aristotle in classical antiquity, and more recently to Charles Darwin, but the association of animals' actions with the actual word "culture" first originated with Japanese primatologists' discoveries of socially-transmitted food behaviours in the 1940s.

What is culture?

Culture can be defined as "all group-typical behavior patterns, shared by members of animal communities, that are to some degree reliant on socially learned and transmitted information".

Organizational culture

One definition of culture, particularly in relation to the organizational aspect is the utilization of "involvement, consistency, adaptation, and mission." Cultural traits that are indicators of a successful form of organization are more likely to be assimilated into our everyday lives. Organizations that utilize the four aforementioned aspects of culture are the ones that are the most successful. Therefore, cultures that are better able to involve their citizens towards a common goal have a much higher rate of effectiveness than those who do not have a shared goal. A further definition of culture is, "[s]ocially transmitted behavior patterns that serve to relate human communities to their ecological settings." This definition connects cultural behavior to the environment. Since culture is a form of adaptation to ones environment, it is mirrored in many aspects of our current and past societies.

Cultural sociology

Other researchers are currently exploring the idea that there is a connection between cultural sociology and psychology. Certain individuals are especially concerned with the analysis of studies connecting "identity, collective memory, social classification, logics of action, and framing." Views of what exactly culture is has been changing due to the recent convergence of sociological and psychological thought on the subject. "Recent work depicts culture as fragmented across groups and inconsistent across its manifestations. The view of culture as values that diffuse other aspects of belief, intention, and collective life has succumbed to one of culture as complex rule-like structures that constitute resources that can be put to strategic use." Culture is specific to region and not just one umbrella definition or concept can truly give us the essence of what culture is. Also referenced is the importance of symbols and rituals as cognitive building blocks for a psychological concept of shared culture.

Memes and cultural transmission

Richard Dawkins argues for the existence of a "unit of cultural transmission" called a meme. This concept of memes has become much more accepted as more extensive research has been done into cultural behaviors. Much as one can inherit genes from each parent, it is suggested that individuals acquire memes through imitating what they observe around them. The more relevant actions (actions that increase ones probability of survival), such as architecture and craftwork are more likely to become prevalent, enabling a culture to form. The idea of memes as following a form of Natural Selection was first presented by Daniel Dennett. It has also been argued by Dennett that memes are responsible for the entirety of human consciousness. He claims that everything that constitutes humanity, such as language and music is a result of memes and the unflinching hold they have on our thought processes.

Evolutionary culture

A closely related concept to memes is the idea of evolutionary culture. The validity of the concept of evolutionary culture has been increasing recently due to the re-evaluation of the term by anthropologists. The broadening scope of evolution from simple genes to more abstract concepts, such as designs and behaviors makes the idea of evolutionary culture more plausible. Evolutionary culture theory is defined as "a theory of cultural phylogeny." The idea that all human culture evolved from one main culture has been presented, citing the interconnectedness of languages as one of his examples has also been presented. There is, however, also the possibility for disparate ancestral cultures, in that the cultures we see today may potentially have stemmed from more than one original culture.

Culture in animals

According to the Webster's dictionary definition of culture, learning and transmission are the two main components of culture, specifically referencing tool making and the ability to acquire behaviors that will enhance one's quality of life. Using this definition it is possible to conclude that animals are just as likely to adapt to cultural behaviors as humans. One of the first signs of culture in early humans was the utilization of tools. Chimpanzees have been observed using tools such as rocks and sticks to obtain better access to food. There are other learned activities that have been exhibited by animals as well. Some examples of these activities that have been shown by varied animals are opening oysters, swimming, washing of food, and unsealing tin lids. This acquisition and sharing of behaviors correlates directly to the existence of memes. It especially reinforces the natural selection component, seeing as these actions employed by animals are all mechanisms for making their lives easier, and therefore longer.

History of animal culture theory

Though the idea of 'culture' in animals has only been around for just over half of a century, scientists have been noting social behaviors of animals for centuries. Aristotle was the first to provide evidence of social learning in the songs of birds. Charles Darwin first attempted to find the existence of imitation in animals when attempting to prove his theory that the human mind had evolved from that of lower beings. Darwin was also the first to suggest what became known as social learning in attempting to explain the transmission of an adaptive pattern of behavior through a population of honey bees.

The vast majority of cultural anthropological research has been done on non-human primates, due to their being closest evolutionarily to humans. In non-primate animals, research tends to be limited, and therefore evidence for culture strongly lacking. However, the subject has become more popular recently, and has prompted the initiation of more research into the area.

Dawkins's meme theory

Evolutionary biologist Richard Dawkins made groundbreaking headway into the field of cultural transmission with his 1976 book entitled The Selfish Gene, which focused heavily on the move to evolution being understood primarily by genetic influence. Dawkins coined the term meme, the primary unit of cultural transmission or imitation, to explain an overarching mechanism of how animal behavior is shared and spread to lead to cultural evolution. The use of the word meme was an intentional phonetic derivation of the similar sounding word "gene", which Dawkins asserts to be the primary unit of selection as it lends itself to pathways of biological evolution:
"We need a name for the new replicator, a noun that conveys the idea of a unit of cultural transmission, or a unit of imitation. 'Mimeme' comes from a suitable Greek root, but I want a monosyllable that sounds a bit like 'gene'. I hope my classicist friends will forgive me if I abbreviate mimeme to meme." (Dawkins)
The analogy between the gene and meme proposed by Dawkins as units of evolutionary biology serves to reinforce the idea that there is a particular pathway of transfer associated with each unit that lends itself either to the evolution of genotypic, phenotypic, and/or behavioral patterns within animal groups. Dawkins asserts that in order for cultural evolution to take place, there needs to be (1) variation within the memes present, (2) the capacity for meme replication between two or more parties, and (3) fitness advantages and/or disadvantages with each meme that lead to the selection or rejection of one meme over another. Likewise, these three criteria in the context of genes are also necessary for genetic evolution. However, with the meme unit, cultural transmission has a distinct feature of being capable of taking place by individuals developing varying interpretations of the meme without exactly "copying" it to pass it on. These interpretations lead to the creation of new memes, which are themselves subject to a cyclic process of selection, rejection, or modification.

Whiten's Culture in Chimpanzees

Andrew Whiten, professor of Evolutionary and Developmental Psychology at the University of St. Andrews, contributed to the greater understanding of cultural transmission with his work on chimpanzees. In Cultural Traditions in Chimpanzees, Whiten created a compilation of results from seven long-term studies totaling 151 years of observation analyzing behavioral patterns in different communities of chimpanzees in Africa (read more about it below). The study expanded the notion that cultural behavior lies beyond linguistic mediation, and can be interpreted to include distinctive socially learned behavior such as stone-handling and sweet potato washing in Japanese macaques. The implications of their findings indicate that chimpanzee behavioral patterns mimic the distinct behavioral variants seen in different human populations in which cultural transmission has generally always been an accepted concept.

Cavalli-Sforza and Feldman models

Population geneticists Cavalli-Sforza & Feldman have also been frontrunners in the field of cultural transmission, describing behavioral "traits" as characteristics pertaining to a culture that are recognizable within that culture. Using a quantifiable approach, Cavalli-Sforza & Feldman were able to produce mathematical models for three forms of cultural transmission, each of which have distinct effects on socialization: vertical, horizontal, and oblique.
  • Vertical transmission occurs from parents to offspring and is a function which shows that the probability that parents of specific types give rise to an offspring of their own or of another type. Vertical transmission, in this sense, is similar to genetic transmission in biological evolution as mathematical models for gene transmission account for variation. Vertical transmission also contributes strongly to the buildup of between-population variation.
  • Horizontal transmission is cultural transmission taking place among peers in a given population. While horizontal transmission is expected to result in faster within-group evolution due to the relationship building between peers of a population, it is expected to result in less between-group variation than the vertical transmission model would allow for.
  • Oblique transmission is cultural transmission being passed from one generation to another younger generation, such as is done by teaching, and the result of reproducing information across generations is a rapid loss of variation within that specific population. Unlike vertical transmission, oblique transmission doesn't need to occur strictly between parent and offspring; it can occur between less-related generations (e.g. from grandparent to grandchild), or from an individual to a non-related younger individual of the same species.

Mechanisms of cultural transmission in animals

Cultural transmission, also known as cultural learning, is the process and method of passing on socially learned information. Within a species, cultural transmission is greatly influenced by how adults socialize with each other and with their young. Differences in cultural transmission across species have been thought to be largely affected by external factors, such as the physical environment, that may lead an individual to interpret a traditional concept in a novel way. The environmental stimuli that contribute to this variance can include climate, migration patterns, conflict, suitability for survival, and endemic pathogens. Cultural transmission can also vary according to different social learning strategies employed at the species and or individual level. Cultural transmission is hypothesized to be a critical process for maintaining behavioral characteristics in both humans and nonhuman animals over time, and its existence relies on innovation, imitation, and communication to create and propagate various aspects of animal behavior seen today. 

Culture, when defined as the transmission of behaviors from one generation to the next, can be transmitted among animals through various methods. The most common of these methods include imitation, teaching, and language. Imitation has been found to be one of the most prevalent modes of cultural transmission in non-human animals, while teaching and language are much less widespread, with the possible exceptions of primates and cetaceans. Recent research has suggested that teaching, as opposed to imitation, may be a characteristic of certain animals who have more advanced cultural capacities, though this is debatable.

The likelihood of larger groups within a species developing and sharing these intra-species traditions with peers and offspring is much higher than that of one individual spreading some aspect of animal behavior to one or more members. This is why cultural transmission has been shown to be superior to individual learning, as it is a more efficient manner of spreading traditions and allowing members of a species to collectively inherit more adaptive behavior. This process by which offspring within a species acquires his or her own culture through mimicry or being introduced to traditions is referred to as enculturation. The role of cultural transmission in cultural evolution, then, is to provide the outlet for which organisms create and spread traditions that shape patterns of animal behavior visibly over generations.

Genetic vs. cultural transmission

Culture, which was once thought of as a uniquely human trait, is now firmly established as a common trait among animals and is not merely a set of related behaviors passed on by genetic transmission as some have argued. Genetic transmission, like cultural transmission, is a means of passing behavioral traits from one individual to another. The main difference is that genetic transmission is the transfer of behavioral traits from one individual to another through genes which are transferred to an organism from its parents during the fertilization of the egg. As can be seen, genetic transmission can only occur once during the lifetime of an organism. Thus, genetic transmission is quite slow compared to the relative speed of cultural transmission. In cultural transmission, behavioral information is passed through means of verbal, visual, or written methods of teaching. Therefore, in cultural transmission, new behaviors can be learned by many organisms in a matter of days and hours rather than the many years of reproduction it would take for a behavior to spread among organisms in genetic transmission.

Social learning

Culture can be transmitted among animals through various methods, the most common of which include imitation, teaching, and language. Imitation is one of the most prevalent modes of cultural transmission in non-human animals, while teaching and language are much less widespread. In a study on food acquisition techniques in meerkats (Suricata suricatta), researchers found evidence that meerkats learned foraging tricks through imitation of conspecifics. The experimental setup consisted of an apparatus containing food with two possible methods that could be used to obtain the food. Naïve meerkats learned and used the method exhibited by the "demonstrator" meerkat trained in one of the two techniques. Although in this case, imitation is not the clear mechanism of learning given that the naïve meerkat could simply have been drawn to certain features of the apparatus from observing the "demonstrator" meerkat and from there discovered the technique on their own.

Teaching

Teaching is often considered one mechanism of social learning, and occurs when knowledgeable individuals of some species have been known to teach others. For this to occur, a teacher must change its behavior when interacting with a naïve individual and incur an initial cost from teaching, while an observer must acquire skills rapidly as a direct consequence. 

Until recently, teaching was a skill that was thought to be uniquely human. Now, as research has increased into the transmission of culture in animals, the role of teaching among animal groups has become apparent. Teaching is not merely limited to mammals either. Many insects, for example have been observed demonstrating various forms of teaching in order to obtain food. Ants, for example, will guide each other to food sources through a process called "tandem running", in which an ant will guide a companion ant to a source of food. It has been suggested that the "pupil" ant is able to learn this route in order to obtain food in the future or teach the route to other ants. There have been various recent studies that show that cetaceans are able to transmit culture through teaching as well. Killer whales are known to "intentionally beach" themselves in order to catch and eat pinnipeds who are breeding on the shore. Mother killer whales teach their young to catch pinnipeds by pushing them onto the shore and encouraging them to attack and eat the prey. Because the mother killer whale is altering her behavior in order to help her offspring learn to catch prey, this is evidence of teaching and cultural learning. The intentional beaching of the killer whales, along with other cetacean behaviors such as the variations of songs among humpback whales and the sponging technique used by the bottlenose dolphin to obtain food, provide substantial support for the idea of cetacean cultural transmission.

Teaching is arguably the social learning mechanism that affords the highest fidelity of information transfer between individuals and generations, and allows a direct pathway through which local traditions can be passed down and transmitted.

Imitation

Imitation can be found in a few members of the avian world, in particular the parrot. Imitation forms the basis of culture, but does not on its own imply culture.

Imitation is often misinterpreted as merely the observation and copying of another's actions. This would be known as mimicry, because the repetition of the observed action is done for no other purpose than to copy the original doer or speaker. In the scientific community, imitation is rather the process in which an organism purposefully observes and copies the methods of another in order to achieve a tangible goal. Therefore, the identification and classification of animal behavior as being imitation has been very difficult. Recent research into imitation in animals has resulted in the tentative labeling of certain species of birds, monkeys, apes, and cetaceans as having the capacity for imitation. For example, a Grey parrot by the name of Alex underwent a series of tests and experiments at the University of Arizona in which scientist Irene Pepperberg judged his ability to imitate the human language in order to create vocalizations and object labels. Through the efforts of Pepperberg, Alex has been able to learn a large vocabulary of English words and phrases. Alex can then combine these words and phrases to make completely new words which are meaningless, but utilize the phonetic rules of the English language. Alex's capabilities of using and understanding more than 80 words, along with his ability to put together short phrases, demonstrates how birds, who many people do not credit with having deep intellect, can actually imitate and use rudimentary language skills in an effective manner. The results of this experiment culminated with the conclusion that the use of the English language to refer to objects is not unique to humans and is arguably true imitation, a basic form of cultural learning found in young children.

Language

Language is another key indicator of animals who have greater potential to possess culture. Though animals do not naturally use words like humans when they are communicating, the well-known parrot Alex demonstrated that even animals with small brains, but are adept at imitation can have a deeper understanding of language after lengthy training. A bonobo named Kanzi has taken the use of the English language even further. Kanzi was taught to recognize words and their associations by using a lexigram board. Through observation of its mother's language training, Kanzi was able to learn how to use the lexigrams to obtain food and other items that he desired. Also, Kanzi is able to use his understanding of lexigrams to decipher and comprehend simple sentences. For example, when he was told to "give the doggie a shot," Kanzi grabbed a toy dog and a syringe and gave it a realistic injection. This type of advanced behavior and comprehension is what scientists have used as evidence for language-based culture in animals.

Primate culture

A bonobo fishing for termites using a sharpened stick. Tool usage in acquiring food is believed to be a cultural behavior.

The beginning of the modern era of animal culture research in the middle of the 20th century came with the gradual acceptance of the term "culture" in referring to animals. Japan's leading primatologist of the time, Kinji Imanishi, first used the word with a prefix as the term "pre-culture" in referring to the now famous potato-washing behavior of Japanese macaques. In 1948, Imanishi and his colleagues began studying macaques across Japan, and began to notice differences among the different groups of primates, both in social patterns and feeding behavior. In one area, paternal care was the social norm, while this behavior was absent elsewhere. One of the groups commonly dug up and ate the tubers and bulbs of several plants, while monkeys from other groups would not even put these in their mouths. Imanishi had reasoned that, "if one defines culture as learned by offspring from parents, then differences in the way of life of members of the same species belonging to different social groups could be attributed to culture." Following this logic, the differences Imanishi and his colleagues observed among the different groups of macaques may suggest that they had arisen as a part of the groups' unique cultures. The most famous of these eating behaviors was observed on the island of Koshima, where one young female was observed carrying soiled sweet potatoes to a small stream, where she proceeded to wash off all of the sand and dirt before eating. This behavior was then observed in one of the monkey's playmates, then her mother and a few other playmates. The potato-washing eventually spread throughout the whole macaque colony, encouraging Imanishi to refer to the behavior as "pre-culture," explaining that, "we must not overestimate the situation and say that 'monkeys have culture' and then confuse it with human culture." At this point, most of the observed behaviors in animals, like those observed by Imanishi, were related to survival in some way.

A chimpanzee mother and baby.

The first evidence of apparently arbitrary traditions came in the late-1970s, also in the behavior of primates. At this time, researchers McGrew and Tutin found a social grooming handclasp behavior to be prevalent in a certain troop of chimpanzees in Tanzania, but not found in other groups nearby. This grooming behavior involved one chimpanzee taking hold of the hand of another and lifting it into the air, allowing the two to groom each other's armpits. Though this would seem to make grooming of the armpits easier, the behavior actually has no apparent advantage. As the primatologist Frans de Waal explains from his later observations of the hand-clasp grooming behavior in a different group of chimpanzees, "A unique property of the handclasp grooming posture is that it is not required for grooming the armpit of another individual... Thus it appears to yield no obvious benefits or rewards to the groomers."

Prior to these findings, opponents to the idea of animal culture had argued that the behaviors being called cultural were simply behaviors that had evolutionarily evolved due to their importance to survival. After the identification of this initial non-evolutionarily advantageous evidence of culture, scientists began to find differences in group behaviors or traditions in various groups of primates, specifically in Africa. More than 40 different populations of wild chimpanzees have been studied across Africa, between which many species-specific, as well as population-specific, behaviors have been observed. The researching scientists found 65 different categories of behaviors among these various groups of chimpanzees, including the use of leaves, sticks, branches, and stones for communication, play, food gathering or eating, and comfort. Each of the groups used the tools slightly differently, and this usage was passed from chimpanzee to chimpanzee within the group through a complex mix of imitation and social learning.

Chimpanzees

In 1999, Whiten et al. examined data from 151 years of chimpanzee observation in an attempt to discover how much cultural variation existed between populations of the species. The synthesis of their studies consisted of two phases, in which they (1) created a comprehensive list of cultural variant behavior specific to certain populations of chimpanzees and (2) rated the behavior as either customary – occurring in all individuals within that population; habitual – not present in all individuals, but repeated in several individuals; present – neither customary or habitual but clearly identified; absent – instance of behavior not recorded and has no ecological explanation; ecological – absence of behavior can be attributed to ecological features or lack thereof in the environment, or of unknown origin. Their results were extensive: of the 65 categories of behavior studied, 39 (including grooming, tool usage and courtship behaviors) were found to be habitual in some communities but nonexistent in others. 

Whiten et al. further made sure that these local traditions were not due to differences in ecology, and defined cultural behaviors as behaviors that are "transmitted repeatedly through social or observational learning to become a population-level characteristic". Eight years later, after "conducting large-scale controlled social-diffusion experiments with captive groups", Whiten et al. stated further that "alternative foraging techniques seeded in different groups of chimpanzees spread differentially...across two further groups with substantial fidelity".

This finding confirms not only that nonhuman species can maintain unique cultural traditions; it also shows that they can pass these traditions on from one population to another. The Whiten articles are a tribute to the unique inventiveness of wild chimpanzees, and help prove that humans' impressive capacity for culture and cultural transmission dates back to the now-extinct common ancestor we share with chimpanzees.

Similar to humans, social structure plays an important role in cultural transmission in chimpanzees. Victoria Horner conducted an experiment where an older, higher ranking individual and a younger, lower ranking individual were both taught the same task with only slight aesthetic modification. She found that chimpanzees tended to imitate the behaviors of the older, higher ranking chimpanzee as opposed to the younger, lower ranking individual when given a choice. It is believed that the older higher ranking individual had gained a level of 'prestige' within the group. This research demonstrates that culturally transmitted behaviors are often learned from individuals that are respected by the group. 

The older, higher ranking individual's success in similar situations in the past led the other individuals to believe that their fitness would be greater by imitating the actions of the successful individual. This shows that not only are chimpanzees imitating behaviors of other individuals, they are choosing which individuals they should imitate in order to increase their own fitness. This type of behavior is very common in human culture as well. People will seek to imitate the behaviors of an individual that has earned respect through their actions. From this information, it is evident that the cultural transmission system of chimpanzees is more complex than previous research would indicate.

Chimpanzees have been known to use tools for as long as they have been studied. Andrew Whiten found that chimpanzees not only use tools, but also conform to using the same method as the majority of individuals in the group. This conformity bias is prevalent in human culture as well and is commonly referred to as peer pressure. 

The results from the research of Victoria Horner and Andrew Whiten show that chimpanzee social structures and human social structures have more similarities than previously thought.

Cetacean culture

Second only to non-human primates, culture in species within the order Cetacea, which includes whales, dolphins, and porpoises, has been studied for numerous years. In these animals, much of the evidence for culture comes from vocalizations and feeding behaviors. 

Cetacean vocalizations have been studied for many years, specifically those of the bottlenose dolphin, humpback whale, killer whale, and sperm whale. Since the early 1970s, scientists have studied these four species in depth, finding potential cultural attributes within group dialects, foraging, and migratory traditions. Hal Whitehead, a leading cetologist, and his colleagues conducted a study in 1992 of sperm whale groups in the South Pacific, finding that groups tended to be clustered based on their vocal dialects. The differences in the whales' songs among and between the various groups could not be explained genetically or ecologically, and thus was attributed to social learning. In mammals such as these sperm whales or bottlenose dolphins, the decision on whether an animal has the capacity for culture comes from more than simple behavioral observations. As described by ecologist Brooke Sergeant, "on the basis of life-history characteristics, social patterns, and ecological environments, bottlenose dolphins have been considered likely candidates for socially learned and cultural behaviors," due to being large-brained and capable of vocal and motor imitation.[35] In dolphins, scientists have focused mostly on foraging and vocal behaviors, though many worry about the fact that social functions for the behaviors have not yet been found. As with primates, many humans are reluctantly willing, yet ever so slightly willing, to accept the notion of cetacean culture, when well evidenced, due to their similarity to humans in having "long lifetimes, advanced cognitive abilities, and prolonged parental care."

Matrilineal whales

In the cases of three species of matrilineal cetaceans, including pilot whales, sperm whales, and killer whales, mitochondrial DNA nucleotide diversities are about ten times lower than other species of whale. Whitehead found that this low mtDNA nucleotide diversity yet high diversity in matrilineal whale culture may be attributed to cultural transmission, since learned cultural traits have the ability to have the same effect as normal maternally inherited mtDNA. The feeding specializations of these toothed whales are proposed to have led to the divergence of the sympatric "resident" and "transient" forms of killer whales off Vancouver Island, in which resident killer whales feed on fish and squid, and transient whales feed on marine mammals. Vocalizations have also been proven to be culturally acquired in killer and sperm whale populations, as evidenced by the distinct vocalization patterns maintained by members of these different species even in cases where more than one species may occupy one home range. Further study is being done in the matrilineal whales to uncover the cultural transmission mechanisms associated with other advanced techniques, such as migration strategies, new foraging techniques, and babysitting.

Dolphins

By using a "process of elimination" approach, researchers Krutzen et al. reported evidence of culturally transmitted tool use in bottlenose dolphins (Tursiops sp.). It has been previously noted that tool use in foraging, called "sponging" exists in this species. "Sponging" describes a behavior where a dolphin will break off a marine sponge, wear it over its rostrum, and use it to probe for fish. Using various genetic techniques, Krutzen et al. showed that the behavior of "sponging" is vertically transmitted from the mother, with most spongers being female. Additionally, they found high levels of genetic relatedness from spongers suggesting recent ancestry and the existence of a phenomenon researchers call a "sponging eve".

In order to make a case for cultural transmission as the mode of behavioral inheritance in this case, Krutzen et al. needed to rule out possible genetic and ecological explanations. Krutzen et al. refer to data that indicate both spongers and nonspongers use the same habitat for foraging. Using mitochondrial DNA data, Krutzen et al. found a significant non-random association between the types of mitochondrial DNA pattern and sponging. Because mitochondrial DNA is inherited maternally, this result suggests sponging is passed from the mother.

In a later study one more possible explanation for the transmission of sponging was ruled out in favor of cultural transmission. Scientists from the same lab looked at the possibility that 1.) the tendency for "sponging" was due to a genetic difference in diving ability and 2.) that these genes were under selection. From a test of 29 spongers and 54 nonspongers, the results showed that the coding mitochondrial genes were not a significant predictor of sponging behavior. Additionally, there was no evidence of selection in the investigated genes.

Rat culture

Notable research has been done with black rats and Norwegian rats. Among studies of rat culture, the most widely discussed research is that performed by Joseph Terkel in 1991 on a species of black rats that he had originally observed in the wild in Palestine. Terkel conducted an in-depth study aimed to determine whether the observed behavior, the systematic stripping of pine cone scales from pine cones prior to eating, was a socially acquired behavior, as this action had not been observed elsewhere. The experimentation with and observation of these black rats was one of the first to integrate field observations with laboratory experiments to analyze the social learning involved. From the combination of these two types of research, Terkel was able to analyze the mechanisms involved in this social learning to determine that this eating behavior resulted from a combination of ecology and cultural transmission, as the rats could not figure out how to eat the pinecones without being "shown" by mature rats. Though this research is fairly recent, it is often used as a prime example of evidence for culture in non-primate, non-cetacean beings. Animal migration may be in part cultural; released ungulates have to learn over generations the seasonal changes in local vegetation.

In the black rat (Rattus rattus), social transmission appears to be the mechanism of how optimal foraging techniques are transmitted. In this habitat, the rats’ only source of food is pine seeds that they obtain from pine cones. Terkel et al. studied the way in which the rats obtained the seeds and the method that this strategy was transmitted to subsequent generations. Terkel et al. found that there was an optimal strategy for obtaining the seeds that minimized energy inputs and maximized outputs. Naïve rats that did not use this strategy could not learn it from trial and error or from watching experienced rats. Only young offspring could learn the technique. Additionally, from cross-fostering experiments where pups of naïve mothers were placed with experienced mothers and vice versa, those pups placed with experienced mothers learned the technique while those with naïve mothers did not. This result suggests that this optimal foraging technique is socially rather than genetically transmitted.

Avian culture

The songs of starlings have been discovered to show regional "dialects," a trait that has potential to have a cultural basis.
Birds have been a strong study subject on the topic of culture due to their observed vocal "dialects" similar to those studied in the cetaceans. These dialects were first discovered by zoologist Peter Marler, who noted the geographic variation in the songs of various songbirds. Many scientists have found that, in attempting to study these animals, they approach a stumbling block in that it is difficult to understand these animals' societies due to their being so different from our own. This makes it difficult to understand the animals' behaviors, let alone determine whether they are cultural or simply practical. 

However, despite this hindrance, evidence for differing dialects among songbird populations has been discovered, especially in sparrows, starlings, and cowbirds. In these birds, scientists have found strong evidence for imitation-based learning, one of the main types of social learning. Though the songbirds obviously learn their songs through imitating other birds, many scientists remain skeptical about the correlation between this and culture: "...the ability to imitate sound may be as reflexive and cognitively uncomplicated as the ability to breathe. It is how imitation affects and is affected by context, by ongoing social behavior, that must be studied before assuming its explanatory power." The scientists have found that simple imitation does not itself lay the ground for culture, whether in humans or birds, but rather it is how this imitation affects the social life of an individual that matters.

Examples of culturally transmitted behaviors in birds

The complexity of several avian behaviors can be explained by the accumulation of cultural traits over many generations.

Bird song

In an experiment regarding at vocal behavior in birds, researchers Marler & Tamura found evidence of song dialects in a sparrow species known as Zonotrichia leucophrys. Located in the eastern and southern parts of North America, these white-crowned song-birds exhibit learned vocal behavior. Marler & Tamura found that while song variation existed between individual birds, each population of birds had a distinct song pattern that varied in accordance to geographical location. For this reason, Marler and Tamura called the patterns of each region a "dialect": however, this term has since been disputed, as different types of in bird song are much less distinct than dialects in human language.

By raising male sparrows in various acoustic settings and observing effects on their verbal behavior, Marler and Tamura found that sparrows learned songs during the first 100 days of their lives. In this experimental setting, male birds in acoustic chambers were exposed to recorded sounds played through a loudspeaker. They also showed that white-crowned sparrows only learn songs recorded from other members of their species. Marler and Tamura noted that this case of cultural transmission was interesting because it required no social bond between the learner and the emitter of sound (since all sounds originated from a loudspeaker in their experiments). However, the presence of social bonds strongly facilitates song imitation in certain songbirds. Zebra finches rarely imitate songs played from a loudspeaker, but they regularly imitate songs of an adult bird after only a few hours of interaction. Interestingly, imitation in zebra finches is inhibited when the number of siblings (pupils) increases.

Innovative foraging

In 20th century Britain, bottled milk was delivered to households in the early morning by milkmen and left on doorsteps to be collected. Birds such as tits (Paridae) began to attack the bottles, opening the foil or cardboard lids and drinking the cream of the top. It was later shown that this innovative behavior arose independently in several different sites and spread horizontally (i.e. between living members) in the existing population. Later experimental evidence showed that conformity may lead to the horizontal spread of innovative behaviors in wild birds, and that this may in turn result in a lasting cultural tradition.

A spread of new foraging behaviors also occurred in an Argentinian population of kelp gulls (Larus dominicanus). During the 20th century, individuals in this population began to non-fatally wound the backs of swimming whales with their beaks, feeding on the blubber and creating deeper lesions in areas that were already wounded. Aerial photographs showed that gull-induced lesions on local whales increased in frequency from 2% to 99% from 1974 to 2011, and that this behavior was not observed in any other kelp gull populations other than two isolated incidents. This implies the emergence and persistence of a local tradition in this population of gulls.

Migration

Juvenile birds that migrate in flocks may learn to navigate accurately through cultural transmission of route choice skills from older birds. Cultural inheritance of migration patterns has been shown in bustards (Otis tarda), and the pattern of inheritance was shown to depend on social structures in the flock.

Avian social networks

Social networks are a specific mechanism of cultural transmission in birds. Information learned in social contexts can allow them to make decisions that lead to increased fitness. A great deal of research has focused on the communication of new foraging locations or behaviors through social networks. These networks are currently being analyzed through computational methods such as network-based diffusion analysis (NBDA).

In wild songbirds, social networks are a mechanism for information transmission both within and between species. Interspecific networks (i.e. networks including birds of different species) were shown to exist in multispecies flocks containing three different types of tits whose niches overlapped. In this study, knowledge about new feeding areas spread through social interactions: more birds visited the new area than the number of birds that discovered the area independently. The researchers noted that information likely travelled faster among members of the same species (conspecifics), but that individuals did not depend solely on conspecifics for transmission. Another study on army-ant-following birds has also evidenced interspecific transmission of foraging information.

A recent study used RFID identification transponders to experimentally manipulate avian social networks: this scanner technology allowed them to restrict access to feeders for some birds and not others. Their data showed that individuals are more likely to learn from those who were able to enter the same feeding area as them. Additionally, the existing "paths" of information transmission were altered following segregation during feeding: this was attributed to changes in the population's social network. 

Others have been able to predict the pattern information transmission among individuals based on a preexisting social network. In this study, social interactions of ravens (Corvus corax) were first analyzed to create a comprehensive network. Then, the order in which individuals learned task-solving behavior from a trained tutor was compared with the network. They not only found that the pattern of learning reflected the network that they had built, but that different types of social connections (such as "affiliative interactions" and "aggressive interactions") characterized different rates of information transmission and observation.

Conformity in avian culture

Bartlett and Slater observed call convergence (i.e. conformity) in budgerigars introduced into groups with different flock-specific calls than their own. They also found that the original calls of flock members did not change significantly during this process.

Conformity is one mechanism through which innovative behaviors can become embedded in culture. In an experimental setting, tits preferentially adopted the locally popular method of opening a two-action puzzle box even after discovering the other possible way of accessing the food. This formed diverging local traditions when different populations were seeded with birds specifically trained in one method.

Other research showed that although conformity has a strong influence on behaviors adopted by birds, the local tradition can be abandoned in favor of an analogous behavior which gives higher reward. This showed that while conformity is a beneficial mechanism for quickly establishing traditions, but that unhelpful traditions will not necessarily be adhered to in the presence of a better alternative.

In some cases, conformity-based aggression may benefit individuals who conform to traditions. Researchers used the framework of sexual selection and conformism in of song types of songbirds to model territorial aggression against individuals with non-conforming song types. Their model showed that aggressors won more frequently when targeting non-conformers (than in un-targeted or random aggression). They also found that alleles for conformity-enforcement propagated more effectively than alleles for tolerance of non-conformity.

Finally, other species of birds have been observed to conform to the personality of other individuals in their presence. Gouldian finches (Erythrura gouldiae) exist in red- and black-headed subtypes, and these subtypes have been shown to have different levels of boldness (measured by the time taken to explore new areas, and other similar tests). Experiments placing black-headed birds (known to be less bold) in the company of red-headed birds (known to be more bold) resulted in the black-headed bird performing "bolder" behaviors, and red-headed birds became "shyer" in the presence of black-headed ones. The experimenters hypothesized that this individual-level conformity could lead to stronger social cohesion.

Fish culture

Guppy mating behavior is believed to be culturally influenced.
 
Evidence for cultural transmission has also been shown in wild fish populations. Scientists Helfman and Schultz conducted translocation experiments with French grunts (Haemulon flavolineatum) where they took fish native to a specific schooling site and transported them to other sites. In this species of fish, the organism uses distinct, traditional migration routes to travel to schooling sites on coral reefs. These routes persisted past one generation and so by relocating the fish to different sites, Helfman and Schultz wanted to see if the new fish could relearn that sites' migration route from the resident fish. Indeed this is what they found: that the newcomers quickly learned the traditional routes and schooling sites. But when residents were removed under similar situations, the new fish did not use the traditional route and instead use new routes, suggesting that the behavior could not be transmitted once the opportunity for learning was no longer there.

In a similar experiment looking at mating sites in blueheaded wrasse (Thalassoma bifasciatum), researcher Warner found that individuals chose mating sites based on social traditions and not based on the resource quality of the site. Warner found that although mating sites were maintained for four generations, when entire local populations were translocated elsewhere, new sites were used and maintained.

Controversies and criticisms

A popular method of approaching the study of animal culture (and its transmission) is the "ethnographic method," which argues that culture causes the geographical differences in the behavioral repertoires of large-brained mammals. However, this significantly downplays the roles that ecology and genetics play in influencing behavioral variation from population to population within a species. Behaviors stemming from genetic or environmental effects are not reliant on socially learned and transmitted information; therefore, they are not cultural.

Culture is just one source of adaptive behavior an organism exhibits to better exploit its environment. When behavioral variation reflects differential phenotypic plasticity, it is due more to ecological pressures than cultural ones. In other words, when an animal changes its behavior over its lifespan, this is most often a result of changes in its environment. Furthermore, animal behavior is also influenced by evolved predispositions, or genetics. It is very possible that "correlation between distance between sites and 'cultural difference' might reflect the well-established correlation between genetic and geographical distances". The farther two populations of a species are separated from each other, the less genetic traits they will share in common, and this may be one source of variance in culture.

Another argument against the "ethnographic method" is that it is impossible to prove that there are absolutely no ecological or genetic factors in any behavior. However, this criticism can also be applied to studies of human culture. Though culture has long been thought to arise and remain independent of genetics, the constraints on the propagation and innovation of cultural techniques inevitably caused by the genome of each respective animal species has led to the theory of gene-culture coevolution, which asserts that "cognitive, affective, and moral capacities" are the product of an evolutionary dynamic involving interactions between genes and culture over extended periods of time. The concept behind gene-culture coevolution is that, though culture plays a huge role in the progression of animal behavior over time, the genes of a particular species have the ability to affect the details of the corresponding culture and its ability to evolve within that species.

We do not know every possible genetic or environmental effect on behavior that exists, nor will we ever. In other words, it is impossible to reject the notion that genes and ecology influence all behaviors, to a degree. Culture can also contribute to differences in behavior, but like genes and environments, it carries different weight in different behaviors. As Laland and Janik explain, "to identify cultural variation, not only is it not sufficient to rule out the possibility that the variation in behavior constitutes unlearned responses to different selection pressures [from the environment], but it is also necessary to consider the possibility of genetic variation precipitating different patterns of learning." Gene-culture coevolution, much like the interaction between cultural transmission and environment, both serve as modifiers to the original theories on cultural transmission and evolution that focused more on differences in the interactions between individuals.

Some scientists believe the study of animal culture should be approached in a different way. Currently, the question being asked is, "is this behavior learned socially (and hence is a result of culture alone), or is it a product of genes and/or environment?" However, it is impossible to find an absolute answer to this question, nor does one in all likelihood exist. Therefore it seems as though scientists should focus on examining how much variance in a behavior can be attributed to culture. Performing field experiments is an excellent way to try to answer this question: translocating individuals between populations or populations between sites could help biologists distinguish between culture, ecology and genetics. For example, if a newly introduced animal shifts its behavior to mimic that of others in its new population, genetic difference as an effect on behavior can be ruled out.

Unanswered questions and future areas of exploration

In the study of social transmissions, one of the important unanswered questions is an explanation of how and why maladaptive social traditions are maintained. For example, in one study on social transmission in guppies (Poecilia reticulata), naïve fish preferred taking a long, energetically costly route to a feeder that they had learned from resident fish rather than take a shorter route. These fish were also slower to learn the new, quicker route compared to naïve fish that had not been trained in the long route. In this case, not only is the social tradition maladaptive, but it also inhibits the acquisition of adaptive behavior.

Inequality (mathematics)

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