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date: 25 April 2024



  • Cedric BoeckxCedric BoeckxICREA (Catalan Institute for Advanced Studies and Research)/Universitat de Barcelona
  •  and Pedro Tiago MartinsPedro Tiago MartinsUniversitat de Barcelona


All humans can acquire at least one natural language. Biolinguistics is the name given to the interdisciplinary enterprise that aims to unveil the biological bases of this unique capacity.


  • Biology of Language

1. History

The idea that the mind is what the brain does, and therefore that language is rooted in the activity of neural cells and assemblies, can of course be traced back to the origin of science itself. In fact, anecdotal evidence concerning the locus of language in the brain can be found in numerous written records. But it is often thought that systematic investigation of the relation between language and the brain (and its evolution) goes back to the discoveries and early theoretical models of Broca and Wernicke. Clearly inspired by F. J. Gall, but also by C. Darwin, these pioneers of neurolinguistics used data from brain lesions to shed light onto the question of brain implementation of complex traits like language.

However, contact between that community and the linguistics community remained scarce. It was not until the beginning of the “cognitive revolution,” under the impetus of Noam Chomsky and Eric Lenneberg, that the seeds of a constructive dialogue between linguistics and the biological sciences began to be sown. Observations concerning language and the brain, or the evolution of language, preceding the cognitive revolution of the mid-1950s ought to be regarded as the pre-history of the field of biolinguistics. The history of the field proper has a much narrower historical span. Some of the early observations belonging to the pre-history of the field may well turn out to be on the right track, but they suffered from a fatal flaw: they failed to combine biological reasoning with a detailed, scientific view of what language is.

It is indeed fair to say that the first sign of a decidedly biological orientation in the study of language, completely foreign at a time when linguistics was dominated by the structuralist paradigm, was Noam Chomsky’s early work in the mid-1950s. Chomsky’s output progressively refined these thoughts, culminating in his Aspects of the Theory of Syntax (1965), whose first chapter articulates the ideas that complement well his review of B. F. Skinner’s Verbal Behavior, in 1959. The latter is generally considered to have contributed decisively to the end of behaviorism, of which Skinner was arguably the fiercest representative. The overarching assumption of Chomsky’s work at this time is that languages are not learned in the conventional sense of the term—say, like a musical instrument, or an academic field—but are rather the product of a biologically determined capacity present in all humans, located in the brain, a great deal of which must be innate, for there is no other way to explain the fact that every speaker produces and understands novel sentences, with no apparent effort and with no explicit instruction. Thus, under this view, which can be assumed as the main tenet of “generative linguistics,” the task of the linguist was taken to be the discovery or formulation of the general principles and rules that generate all possible sentences, and that must nonetheless be restrictive enough so as to not generate impossible sentences. In the event that all of these rules were formulated, their virtual output would amount to a list of all (and only) sentences in a given language. But the core, defining questions of Chomskyan linguistics came from somewhere else: animal behavior.

The following questions, formulated by ethologist Niko Tinbergen in 1963, are known to most linguists familiar with Chomsky’s work:

What stimulates the animal to respond with the behavior it displays, and what are the response mechanisms?

How does an organism develop as the individual matures?

Why is it necessary for the animal’s success and how does evolution act on that behavior?

These questions are at the basis of Chomsky’s own questions about language; they are the template upon which questions about the biology of any behavioral property of organisms should rest, and language is one of them. Chomsky had already recognized the importance of Tinbergen’s insights in his early writings (see Chomsky, 1959, footnotes 1, 31). Thus, the following questions, as summarized by Boeckx and Grohmann (2007, p. 1) became the central problems introduced by Chomskyan linguistics:

What is knowledge of language?

How is that knowledge acquired?

How is that knowledge put to use?

How is that knowledge implemented in the brain?

How did that knowledge emerge in the species?

Some of these questions seem to be answerable be making use of the methods and techniques available to linguistics, but halfway through the list the biological overtones are stronger; they require biology proper.

While Chomsky was laying out the conceptual arguments for a biological orientation in the study of language, which he derived from his study of linguistic structures and sound logic, Lenneberg was exploring the strictly biological issues that would remain relevant for decades to come, amassing an impressive body of converging evidence from many fields, which he presented in a very coherent fashion in his Biological Foundations of Language (1967), to which Chomsky also contributed a chapter. Lenneberg offered concrete research directions for the study of language, much different from—but complementary to—Chomsky’s own formal, conceptual work. This work focuses on different aspects of human biology that putatively contribute to language, and which are still very much still relevant today, though some of them need to be re-interpreted in light of current evidence and knowledge. Many of the topics presented by Lenneberg seem to make more sense in today’s evolutionary biology than they did at the time, when the Modern Synthesis still seemed to be all that was needed for understanding all there is to know about the evolution of organisms and their properties. Lenneberg felt that there had to be more than genes at play, for while they are obviously essential, and while genomes are associated with specific organisms, genes do not store traits (or “faculties”) in the way that linguistics—including Chomsky(ans)—sometimes seem to imply. The geno-centric argument makes sense to a certain extent on conceptual grounds, to help tear down barriers and get close to core problems, but Lenneberg’s will to push it further forced him to take the biology of language further as well. Lenneberg is now considered an essential figure in the biolinguistic enterprise because the means now exist to realize that he was right about a lot of things, and that the biology of his time was not adequate. “It is not strictly correct to speak of genes for long ears, for auditory acuity, or for the capacity for language […] [g]enes can only affect ontogenesis through varying the cells’ repertoire of differentiation, but this, in turn, may have secondary effects upon structure, function, and capacities” (Lenneberg, 1967, p. 241). (For a discussion on the relevance of Lenneberg’s views for modern biology, see Boeckx & Longa, 2011.)

The next checkpoint for biolinguistics was a meeting held in 1974, organized by Massimo-Piattelli Palmarini, which brought together a number of researchers in anthropology, biology, linguistics, and other fields, engaging in discussions about language, biology, and the brain that expanded on the work that had been developed up until that point. This meeting also marked the coining of the term “Biolinguistics,” which was chosen to name the enterprise (at first, interchangeably with “bioanthropology”) that appeared to emerge out of the discussions (Piattelli-Palmarini, 1974; Luria, 1974). The term had been used before (e.g., Meader & Muyskens, 1950), but most likely this was not known by Piattelli-Palmarini at the time. Another important meeting took place in 1975 in Paris, between Noam Chomsky and Jean Piaget. This was perhaps an early sign of the importance of eclecticism in biolinguistic discussions, which tend to include perspectives considered to be incompatible, only to uncover points of commonality that the different factions do not usually explore, sometimes for unclear or strictly historical reasons (see Boeckx, 2014a for discussion).

A growing number of scientists were starting to notice that it was indeed possible to look at language differently, outside of the humanities department. There was even talk of a specialized journal, an idea that gained support from people like Noam Chomsky, François Jacob, and Konrad Lorenz, among others, but it never came to fruition (see Boeckx & Grohmann, 2007 for documentation).

The following years were for several reasons important for the development of biolinguistics, but progress has been slow, for different reasons.

2. Limited Impact

Even though support for biolinguistics initially came from both the linguistics and the biology sides, progress has been slow since Chomsky’s and Lenneberg’s earliest writings. Biolinguistics has in practice been seen as a sub-field or rebranding of generative linguistics, and as such most of the work said to be biolinguistic came from there. But the biology is rarely taken seriously into account in such work: even though linguistics managed to incorporate biology into its rhetoric, it has remained a largely descriptive field, with rare signs of real biological work. A brief, random survey of the generativist literature will reveal mentions of biology, but usually confined to introductory sections and final remarks, and never at the center of the discussions, and never too relevant for the work that unfolds. Some would probably counter this claim, and claim instead that they are interested in the biological properties of language, but the very fact that the paragraphs on the biology of language are for the most part interchangeable across papers, frameworks and even eras shows that not a lot of new insights have come out of linguistics since Chomsky’s and Lenneberg’s original ideas, and that the sources of the mentions of biology are not biology papers, which not surprisingly look a little different today than they did in the mid-20th century. The effects of referring to the biological properties of language in a linguistics paper and leaving the subject unaddressed can be seen as quite harmless, but then the question arises of where these properties are to be pursued instead, if at all. The answer is obviously biology, and it motivates the generative linguistics/biolinguistics distinction, as the former does not directly engage with it.

The success of linguistics proper has likely contributed to the relegation of biological concerns to the background. Each generative model became better than the previous one at describing and offering theories of linguistic structure, namely by getting a handle on linguistic variation and its limits. But the core, biolinguistic questions are still there, and linguistics can offer no better answers than it did when they were first formulated. The recent collaboration between prominent generative linguists and non-linguists (case in point: Hauser, Chomsky, & Fitch, 2002) could be a step in the right direction, because collaboration was also at the heart of biolinguistic thought half a century ago, but most of such works still hinge too much on linguistics, and not enough on biology.

One of the reasons why the success of linguistics makes the success of biolinguistics slower is the kind of constructs posited by the former. Across different linguistic frameworks, the postulation of primitive units that must somehow be encoded in Universal Grammar (UG) is common practice. Its goal is to explain grammatical phenomena and in a more general sense the nature of the faculty of language. The field of phonology is perhaps where explanations of this kind find their roots, after the notion of “feature” was introduced, usually attributed to Trubetzkoy. This approach is of course also the one practiced in the domains of morphology and syntax. There are different kinds of features that are taken to be encoded in UG and thus account for the outputs observed. For example, the cartographic enterprise attempts to ground the notion of feature in substance by adopting an empirical perspective: it attempts to explore the “fine structure of the [sentential] left periphery,” and puts forth a highly articulated structure of functional and lexical heads (Rizzi, 1997)—but also morphemes, certain types of adverbs and adjectives, quantifiers, classifiers, numerals, etc. Recent studies in cartography roughly estimate the number of such projections to be up to 400 (Cinque & Rizzi, 2008, p. 47). What accounts for the rising number is the fact that when discussing different grammars, different, “new” features keep “emerging” as explanations of how cross-linguistic differences are built. Even the most minimal morphosyntactic difference can potentially be treated as a sufficient basis for extending one’s chronically incomplete inventory of projections by adding yet another element to it. Yet, despite the pervasiveness of features in all domains of linguistic inquiry, it is dubious whether features can go beyond language-specific particularities and unveil the properties of language as a biological organ, instead of resulting in descriptive grammars of the language (see, among others, Boeckx, 2011, 2014b). There is a very important, independent reason why the primitives posited in linguistic theories might be slowly losing their explanatory power, which has much to do with the confrontation with neighboring fields. If something like [−syllabic] or [+accusative] is given to a neurologist, who according to linguistics should be able to find it in the brain in some relevant manner, the neurologist does not know what to do. This is not due to the limitations of current techniques or technology, but rather to the fact that linguistics and neuroscience do not work with units of the same kind. Similarly, if an evolutionary biologist is asked to trace back the origin of verbal agreement, not much is to be expected. A defense of linguistic primitives could be that each field is expected to work within specific assumptions, methodologies, and units, but if linguistics is to unveil the biological properties of language, the units it proposes should be interpretable by other relevant fields. To our eyes, only a few linguists, such as Ray Jackendoff (see Jackendoff, 2002), have sought to wrestle with this issue, and no clear results can be discerned yet.

Poeppel and Embick (2005) illustrate this problem by comparing the kinds of primitive units usually found in linguistics and the ones found in neuroscience. In linguistics, common primitives include distinctive features, syllables, morphemes, noun phrases, or clauses, while neuroscience features notions like dendrites, spines, neurons, cell-assemblies, population, or cortical column. In the case of operations, linguistics constructs theories of concatenation, linearization, phrase-structure generation, or semantic composition, while neuroscience studies receptive fields, oscillations, and synchronization. Poeppel and Embick argue that there is currently no way to come up with linking hypotheses that bridge the kinds of elements that each of the fields recognizes, and because of that it is simply not possible to ground linguistic units posited thus far in biological terms. They identify two problems that ought to be resolved if this predicament is to be changed: the Granularity Mismatch Problem (GMP) and the Ontological Incommensurability Problem (OIP), defined as follows:

GMP: Linguistic and neuroscientific studies of language operate with objects of different granularity. In particular, linguistic computation involves a number of fine-grained distinctions and explicit computational operations. Neuroscientific approaches to language operate in terms of broader conceptual distinctions.

OIP: The units of linguistic computation and the units of neurological computation are incommensurable.

These two, related, problems concern, respectively, the different nature of primitives in both fields and the fact that they have been developed independently of each other. The result is, indeed, the questionable progress in cognitive neuroscience, more specifically in relation to language. This is a very good illustration of the advantages of true interdisciplinarity: the earlier it starts, the better, for letting fields fully develop the conceptual foundations on which they rest and only later “marrying” them might make the relationship very difficult or impossible, as fields diverge from the continuum of which they should be part. The sui generis nature of linguistics and the primitives it posits might have contributed to its success as a circumscribed field of inquiry, but as long as it continues this way it will not be able to learn from or teach anything to fields that it should have as allies.

3. Reemergence

While it is true that biolinguistics has remained an obscure or largely rhetorical field during the last half century or so, it is also true that the early 21st century has spawned an unprecedented number of publications, conferences, and other initiatives that highlight the importance of biology for the study of language. This has not been by chance, for there are a number of factors that can explain why this has happened only at the turn of the century.

A very important factor is the genomic revolution. After years of discussion on the genetic basis of language—or, rather, about the fact that such a basis must exist—news finally came from genetics. It was discovered that a language disorder that plagues half of the members of an English family is linked to a specific mutation in a gene, the now well-known FOXP2 (Fisher et al., 1998; Lai et al., 2001). The linguistic and cognitive science circles rejoiced; it seemed like the argument was right all along, and that finally the (or at least one) “language gene” had been found, a claim widely reported in the media. Reactions of various kinds did not take long. Soon the gene was built into evolutionary explanations of language, with claims about language being present in our close relatives that too had the modern, “human” version of the gene (e.g., Krause et al., 2007; Reich et al., 2010). Proposing such evolutionary stories requires big leaps of faith, for in reality genes alone don’t guarantee the traits they are associated with, and not only that, but as Benitez-Burraco (2009) reports, there may be more than one hundred genes somehow involved in language. It was discovered shortly after that FOXP2 is a highly conserved gene, which has led to research on its functioning in other, especially vocal-learning species, and on the factors that have contributed to the mutated FOXP2 found in humans. More so than arriving at consensual conclusions on the history and role of FOXP2 and other genes it regulates in language, their study has led to the acceptation of the comparative method so patent in the biological sciences. It also highlighted the bumpy roads between genotypes and phenotypes. In fact, it is now known that very different phenotypes can be originated by the same genotype if the latter is differently affected by the environment (West-Eberhard, 2003). This has great implications for generative linguistics: it means that Universal Grammar or the faculty of language cannot be conceived of as the “linguistic genotype,” in that there is no intrinsic, separate genotype of language. That is to say, modular conceptions of cognitive domains like language are likely to dissolve as we learn more about the (generic) mechanisms implementing cognition at the molecular and cellular levels.

Hauser et al. (2002) introduced the “Faculty of Language—Broad Sense” (FLB)/“Faculty of Language—Narrow Sense” (FLN) distinction, in an attempt to come to grips with the fact language had to evolve somehow, but that something about it is truly unique. FLB is defined as all that enters into the Faculty of Language, most components of which will be found across domains and species, with FLN being the subset of components of FL that are to be found only in humans. Most research has focused on FLN, following tradition (i.e., that there is something biologically unique to human language), but the fact that the distinction was proposed, that is, the acknowledgement that at least part of the language faculty could be specific neither to language nor to humans, represents a great departure from that tradition, and opens ways for the study of animal cognition in service of the study of language. Perhaps the way to best interpret its message is to follow the emerging, bottom-up approach to cognitive psychology, which Waal and Ferrari (2010) herald: “What if we were to replace our obsession with complex cognition with an exploration of basic processes? Instead of asking which species can do X, the question would become how does X actually work?”

Another important factor is the rise of “Evo-Devo” (evolutionary development), which is in effect a new paradigm in biology.

For half a century, biology had been dominated by a genocentric, selectionist framework called the Modern Synthesis, as a result of around 150 years of biological thinking. Before Darwin, there were two major biological questions for which answers had never been provided. The first was how to explain the overwhelming variety and history of life forms in the planet. Species are so different among them that any relation between them seems hard to conceive. The second, related question, was how to account for the fact that form and function seem to go hand-in-hand; all species seem to be perfectly equipped for the tasks they have to carry out in order to survive, from their appearance to the way they move or the way they take in and process the stimuli that are relevant for them. Darwin was the first to develop scientific answers to these questions. Darwin observed that traits vary within populations and are inherited from one generation to the next one—although the exact mechanisms that allowed for this were a mystery to him—and also that certain members of a species will succumb to the environment, while others will survive and prosper. The principles that he derived from these observations provided satisfactory explanations for the two basic biological questions. The first was “descent with modification,” which explains the variety and history of organisms, and the second was “natural selection,” which explains why form and function go hand-in-hand even though nature does not have intentions, and as such cannot be goal-oriented. Later, Mendel’s work on genetics (1866) had its breakthrough, nearly 40 years after its inception. The greatest contribution to the Mendelian revival was perhaps a seminal paper by Ronald Fisher (1918), in which he managed to show, contrary to the opinion at the time, that gradualist character of evolution could still make sense in the context of Mendelian genetics, whose saltational mechanisms had been seen up until that point as problematic for the Darwinists (Box, 1978). After the impetus of Fisher’s individual work, he and colleagues developed the field of population genetics (Fisher, 1918, 1930; Haldane, 1932, 1934; Wright, 1931, 1932, 1937), which studies the frequency and interaction of alleles and genes at the level of populations, thus marking the abandonment of essentialist theories of species-fixation, an idea further developed by Mayr (1942). Further support was provided by Simpson (1944), who argued that population genetics was very much in line with the conclusions and implications of geology and paleontology, whose empirical findings were obviously of great importance to the validation of evolutionary theories.

Roughly, the conjunction of all these elements led to the Modern Synthesis, the body of ideas deemed essential to the understanding of how evolution works, characterized by genocentricity. This was the dominating paradigm at the time of the cognitive revolution of the 1950s, and most of its rhetoric remained in linguistics for decades, almost invariably resulting in genocentric accounts of language.

The advent of Evo-Devo brought a different, more inclusive perspective to biology, in that it showed that the tenets of the Modern Synthesis were limited, and brought in considerations from a host of fields that move genocentric and selectionist accounts from the center, and give equal—if not more—importance to the environment and development. This introduced a pluralistic, seriously interdisciplinary agenda of inquiry that is nowadays at the heart of the biolinguistic approach. This state of affairs lends itself to the exploration of language in new ways, and it is expected to allow the formulation of questions and hypotheses that go very much beyond a genetically determined, fixed faculty of language. But it is still early days.

The genomic revolution made the limits of genocentrism patently obvious, and gradually displaced selectionist considerations from the center to the periphery, opening up new lines of inquiry animated by a more eclectic, pluralistic agenda. This agenda is no longer a single-level model, with genes at its central causal force, but a much more interactionist perspective with equal weight placed on genes, the environment, and development. The new paradigm has been tentatively called the Extended Synthesis (see Pigliucci & Muller, 2010).

One last main factor, more familiar to linguists—sometimes to the point where it is conflated with biolinguistics—is the advent of linguistic minimalism. This too represents a paradigm shift of sorts in linguistic theory. Linguistic minimalism is the attempt to reduce the specifically linguistic system to the minimum, relying instead on general cognitive mechanisms for the generation of the rich structure once attributed to a complex, overspecified “Universal Grammar.” Such an approach makes way for the reconciling of linguistic theory with modern biological theory and discoveries, in that it meshes well with the advancements previously discussed. The kinds of operations proposed in minimalist accounts of language lend themselves more easily to explorations of brain implementation, in that they are computationally more plausible as far as the brain is concerned, and can thus be investigated biologically. Another important aspect of minimalism is the recognition that what were once taboos in generativism, namely social transmission, cultural evolution, or more generally the environment, are important factors in the emergence of grammatical properties. Ignoring these factors was perhaps important conceptually, as a way of emphasizing the biological character of language, but now we are at the point when we know that biology is more than a genetic blueprint. This makes possible the rapprochement of ideas traditionally shunned by the generativist community for historical reasons. In a sense, the very success of the minimalist program is contingent on the success of external, “non-UG” factors, and by extension on the abandonment of genocentric accounts of language that were at the heart of early biolinguistic thought.

4. Future

Biolinguistics is better characterized as an approach, rather than a field. This is because one of the goals of biolinguistics is very much a methodological one: to bring together insights and discoveries from various fields productively. To achieve this, more than vague acquaintance with the fields mentioned thus far is necessary. A biolinguist must be well-versed in the biological literature, and must seek collaboration with other researchers who are experts on biological sub-fields. It is important to resist vague mentions of other fields without really exploring the issues they are concerned with; the latter is more likely to alienate those fields, as is so often the case. This endeavor must also rely on encouragement and resources made available to young researchers who are interested in the biology of language, from exposure to different perspectives to the culture of the fields in the life sciences that offer them, which function very differently from linguistics (e.g., reading or publishing a biology paper is quite a different experience when compared to a linguistics paper). If research takes the biolinguist there, the biolinguist should not be surprised to study something that apparently has little to do with language as traditionally conceived, and should welcome the study of other domains and other species entirely (e.g., birdsong or primate tool use). This does not mean coming up with analogies of what those species are doing and language (as is common in media reports, but also in the technical literature). Instead, it means decomposing and making sense of language in an evolutionary context. Doing so will reinvigorate the work of a researcher of language: it opens up immense possibilities of what can be studied, where one can go to study it, and where it can be shared with the scientific community.

A readily available avenue for research lies in the bottom-up approach, which brings together modern comparative cognition and minimalism. Going this route requires stretching theories of computation to the point where they can make sense in neurological and genetic terms, and not merely serving as formal accounts of what could be happening in a computer if the mind worked exactly like one. To get to the point where this can be achieved, the role of other factors in the shaping of the language faculty and linguistic structure must be taken into account, and the still evolving evolutionary biology provides a good basis for that exploration, and brings the study of language within the fold of an Extended Synthesis. The key is bringing knowledge together through linking hypothesis, and for the hypothesis to link, the different elements must have things in common. There are two ways of looking at this challenge. The first is the point of view of the non-linguists, who refer to language and use language in their work without necessarily knowing what language is, and different working definitions abound in the literature. This is not to say that all linguists are clear on what language is—they are not—but others tend to look at it simplistically without ever having studied it.

The other way of looking at this challenge is the way in which linguists have made their results available (which might have a strong influence on the previous one). Psychologists, neuroscientists, and geneticists are used to understanding what their colleagues mean, even if it takes some familiarization, but when they look into linguistics they are likely to find what seems to be cryptic and circumscribed terminology and ideas that they can simply not exploit. The result is often either ignoring the work of linguists or misinterpreting it. Some linguists who are keen on the biological nature of language have tried to (re)formulate their insights in such a way that cross-discipline discussion is possible, but this is very much an ongoing and embryonic task.

There is a need to take linguistic theory and filter out computational primitives not in service of the theory itself, but instead in service of what they are supposed to account for, namely neuronal computation. For a linguistic model to be tackled beyond linguistics, it must offer a set of essential primitives that abide by principles of computation at a fine enough grain, that is, whose brain implementation can be fathomed and not just assumed. Doing so will provide the pieces that make it possible to explore the evolution and nature of language in the big picture: its potentially distant origins, how they got together, and how they are implemented neuronally. This would be a step toward solving the aforementioned Granularity Mismatch Problem that Poeppel and Embick (2005) bring to attention. The other problem they highlight, the Ontological Incommensurability Problem, is a more serious one, which could be true in principle. But it should not be taken as an irreversible fact. Instead, it should be taken as something that can be remedied if different scientists go the extra mile and adopt a vertical, multilevel approach, not unlike what Marr (1982) proposed for vision (linking the computational, algorithmic, and implementational levels of analysis). The stale character of ontological incommensurable units can be replaced if they are decomposed and their relations explored. Linking specific brain areas to language or specific linguistic operations only contributes to their imperviousness to investigation, and it will be more fruitful to look for basic computation that is recruited for higher-level cognitive functions than what we call language is a manifestation of. Mapping specific functions to specific brain areas—methodological assumptions aside—will give us information about only where in the brain something is happening, and not what that is or how it is accomplished. This is an assumption that not only linguists but also brain researchers are still working on surpassing (see Poeppel, 2012).

An old idea that still plagues biological inquiry but that must be abandoned is that the faculty of language is uniform across individuals. Again, while this is important when delineating the object of study, the point has come where evidence to the contrary is available. The fact that different modalities coexist, for example, is good evidence. The study of linguistic disorders also allows us look at them as breakdowns of the language faculty, and these are widely varied. Also, when taking the neurological level into account, language acquisition is not as straightforwardly universal as once thought. Similar milestones are still cleared across individuals, but they also vary. The same way that different phenotypes can result from the genotypes, similar cognitive profiles can also rely on different brain architectures. The dynamic interaction of all the different factors that make up biological structures and behavior—encompassed by evolution, development, and the environment—result in different ways of implementing a functional faculty such as language.

After five decades of biolinguistics, a point has been reached where testable hypotheses about the nature and evolution of language can be formulated. Language has become a subject not only of linguistics, but crucially of a number of different disciplines as well, working in tandem. Much work needs to be done, and methodological barriers pose serious challenges, but as researchers progressively learn to bring these disciplines together, the study of language becomes a much larger enterprise, and new avenues of research open up. At the same time, uncovering the roles of hitherto-ignored factors in the shaping of the language faculty results in the reduction of what is to be explained, bringing us closer to the goal of discovering what is unique about our species.

Further Reading

  • Amundson, R. (2005). The changing role of the embryo in evolutionary thought: Roots of evo-devo. Cambridge, U.K., and New York: Cambridge University Press.
  • Boeckx, C. (2010). Language in cognition: Uncovering mental structures and the rules behind them. Malden, Mass.: Wiley-Blackwell.
  • Boeckx, C., & Grohmann, K. (Eds.). (2013). The Cambridge handbook of biolinguistics. Cambridge, UK: Cambridge University Press.
  • Buszáki, G. (2006). Rhythms of the brain. Oxford: Oxford University Press.
  • Chomsky, N. (1980). Rules and representations. New York: Columbia University Press.
  • Dediu, D. (2015). An Introduction to genetics for language scientists. Cambridge, U.K.: Cambridge University Press.
  • Di Sciullo, A. M., & Boeckx, C. (2011). The biolinguistic enterprise: New perspectives on the evolution and nature of the human language faculty. Oxford: Oxford University Press.
  • Jenkins, L. (2000). Biolingistics. Cambridge, U.K.: Cambridge University Press.
  • Lenneberg, E. H. (1967). Biological foundations of language. New York: Wiley.
  • Marcus, G. (2004). The birth of the mind. New York: Basic Books.
  • Pigliucci, M., & Müller, G. B. (2010). Evolution: The extended synthesis. Cambridge, Mass.: MIT Press.


  • Benitez-Burraco, A. (2009). Genes y lenguaje, aspectos ontogenéticos, filogenéticos y cognitivos. Barcelona: Reverté.
  • Boeckx, C. (2011). Review of Anna Kibort & Greville G. Corbett (Eds.), Features: Perspectives on a key notion in linguistics (Oxford: Oxford University Press, 2010); Journal of Linguistics, 47(2), 522–524.
  • Boeckx, C. (2014a). The roots of current biolinguistic thought: Revisiting the “Chomsky-Piaget” debate in the context of the revival of biolinguistics. Teorema: Revista internacional de filosofía, 33(1), 83–94.
  • Boeckx, C. (2014b). Elementary syntactic structures. Cambridge, U.K.: Cambridge University Press.
  • Boeckx, C., & Grohmann, K. (2007). The Biolinguistics manifesto. Biolinguistics, 1(1), 1–8.
  • Boeckx, C., & Longa, V. M. (2011). Lenneberg’s views on language development and evolution and their relevance for modern biolinguistics. Biolinguistics, 5(3), 254–273.
  • Box, J. F. (1978). R. A. Fisher: The life of a scientist. New York: Wiley.
  • Chomsky, N. (1959). A review of B. F. Skinner’s Verbal Behavior. Language, 35(1), 26–58.
  • Chomsky, N. (1965). Aspects of the theory of syntax. Cambridge, Mass.: MIT Press.
  • Cinque, G., & Rizzi, L. (2008). The cartography of syntactic structures. In V. Moscati (Ed.), CISCL working papers on language and cognition, Vol. 2. (pp. 43–59). Siena, Italy: CISCL Università di Siena.
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