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Stephen R. Anderson
Yale University
[Prepared for Cambridge
Encyclopedia of the Linguistic Sciences]

An understanding of the communicative capacities of other animals is important

on its face both for an appreciation of the place of human language in a broader
context, and also as a prerequisite to discussion of the evolution of language. On
closer examination, however, the differences between human language and the
systems of other animals appear so profound as to make both projects more
problematic than they appear at first.
In the 1950s and 1960s, ethologists like Konrad Lorenz and Niko Tinbergen
revolutionized behavioral biologists views of the cognitive capacities of animals,
but consideration of animal communication focused on the properties of quite
simple systems. A prime example of communication in the texts of the time was
the stickleback. A crucial component of the mating behavior of this common fish is
the pronounced red coloration of the males under-belly when he is in mating
condition, which furnishes a signal to the female that she should follow him to his
pre-constructed nest, where her eggs will be fertilized. On this model,
communication was viewed as behavioral or other signals emitted by one
organism, from which another organism (typically, though not always, a
conspecific) derives some information. The biological analysis of communication
thus came to be the study of the ways in which such simple signals arise in the
behavioral repertoire of animals and come to play the roles they do for others who
perceive them. Those discussions make little if any contact with the analysis of
human language.
In the intervening half century, we have come to know vastly more about the
nature and architecture of the human language faculty, and to have good reason to
think that much of it is grounded in human biology. One might expect, therefore, to
find these concerns reflected in the behavioral biology literature. A comprehensive
modern textbook on animal communication within this field (e.g. Bradbury &
Vehrenkamp 1998) reveals a great deal about the communicative behavior of
many species and its origins, but within essentially the same picture of what
constitutes communication in (non-human) animals, confined to unitary signals
holistically transmitted and interpreted. Little if any of what we have come to
know about human linguistic communication finds a place here.
Biologists have not in general paid much attention to the specifics of linguistic
research (though their attention has been caught by the notion that human

language is importantly based in human biology), and are often not as

sophisticated as one might wish about the complexity of natural language. But the
consequences of this may not be as serious as linguists are inclined to think: in fact,
the communicative behavior of non-humans in general is essentially encompassed
within the simple signal-passing model. The complexities of structure displayed by
human language are apparently quite unique to our species, and may not be
directly relevant to the analysis of animal communication elsewhere.

What (Other) Animals Do

Communication in the sense of emission and reception of informative signals is
found in animals as simple as bacteria (quorum sensing). Most familiar, perhaps,
are visual displays of various sorts indicating aggression, submission, invitations to
mate, etc. In some instances these may involve quite complex sequences of
gestures, reciprocal interactions, and the like, as in the case of the nesting and
mating behavior of many birds. In others, a simple facial expression, posture, or
manner of walking may provide the signal from which others can derive
information about the animals intentions and attitudes.
These differences of internal structure are of course crucial for the correct
expression and interpretation of a particular signal, but they play little or no role in
determining its meaning. That is, the individual components of the signal do not in
themselves correspond to parts of its meaning, in the sense that varying one subpart results in a corresponding variation in what is signaled. Animal signals,
however complex in form (and however elaborate the message conveyed), are
unitary wholes. An entire courtship dance, perhaps extending over several minutes
or even longer, conveys the sense I am interested in mating with you, providing a
nesting place and care for our offspring. No part of the dance corresponds exactly
to the providing care part of the message; the message cannot be minimally
altered to convey I am interested in mating, but not in providing care for our
offspring, I was interested in mating (but am no longer)... etc. Variations in
intensity of expression can convey (continuous) variations in the intensity of the
message (e.g., urgency of aggressive intent), but that is essentially the only way
messages can be modulated.
The most widely discussed apparent exception to this generalization is the dance
language of some species of honeybees. The bees dance conveys information
about (a) the direction, (b) the distance, and (c) the quality of a food source (or
potential hive site), all on quasi-continuous scales and each in terms of a distinct
dimension of the dance. Although the content of the message here can be
decomposed, and each part associated with a distinct component of the form of the
signal, there is no element of free combination. Every dance necessarily conveys
exactly these three things, and it is only the relative value on each dimension that
is variable. As such, the degree of freedom available to construct new messages is

not interestingly different from that involved in conveying different degrees of fear
or aggression by varying degrees of piloerection.
Visual displays do not at all exhaust the modalities in which animal communication
takes place, of course. Auditory signals are important to many species, including
such classics of the animal communication literature as frog croaks and the calls
and songs of birds. In some species, portions of the auditory spectrum that are
inaccessible to humans are involved, as in the ultrasound communication of bats,
some rodents, and dolphins, and the infrasound signals of elephants. Chemical or
olfactory communication is central to the lives of many animals, including moths,
mice and lemurs as well our pet cats and dogs. More exotic possibilities include the
modulation of electric fields generated (and perceived) by certain species of fish.
In some of these systems the internal structure of the signal may be quite complex,
as in the songs of many oscine songbirds, but the general point made above still
holds: however elaborate its internal form, the signal has a unitary and holistic
relation to the message it conveys. In no case is it possible to construct novel
messages freely by substitutions or other ways of varying aspects of the signals
In most animals, the relation of communicative behavior to the basic biology of the
species is very direct. Perceptual systems are often quite precisely attuned to
signals produced by conspecifics. Thus, the frogs auditory system involves two
separate structures (the amphibian papilla and the basilar papilla) that are
sensitive to acoustic signals, typically at distinct frequencies. The frequencies to
which they are most sensitive vary across species, but are generally closely related
to two regions of prominence in the acoustic structure of that species calls. Mice
(and many other mammals) have two distinct olfactory organs, projecting to quite
distinct parts of the mouse brain. The olfactory epithelium is responsive to a wide
array of smells, but the vomeronasal organ is sensitive exclusively to the
pheremones that play a major role in communication and social organization. In
this case, as in many, many others, the perceptual system is matched to production
in ways that optimize the organisms sensitivity to signals that play a crucial
ecological role in the life of the animal.
The essential connection between a species system of communication and its
biology is also manifested in the fact that nearly all such systems are innately
specified. That is, the ability to produce and interpret relevant signals emerges in
the individual without any necessary role of experience. Animal communication is
not learned (or taught), but rather develops (in the absence of specific pathology,
such as deafness) as part of the normal course of maturation. Animals raised under
conditions in which they are deprived of exposure to normal conspecific behavior
will nonetheless communicate in the fashion normal to their species when given a

Exceptions to this generalization are extremely rare, apart from human language.
Vocal learning, in particular, has been demonstrated only to a limited extent in
cetaceans and some bats, and more extensively in three of the twenty seven orders
of birds. The study of birds, especially oscine songbirds, is particularly instructive
in this regard. In general, their song is learned on the basis of early exposure to
appropriate models, from which they in turn compose their own songs. There is
much variation across species, but a clear generalization emerges: for each species,
there is a specific range of song structures that individuals can learn. Experience
plays a role in providing the models on which adult song is based, but (with the
exception of a few very general mimics, such as the lyrebird) this role is quite
narrowly constrained by the song-learning system of the individual species.

What Humans Do, and How it is Different

Like the systems of communication of other animals, human language is deeply
embedded in human biology. Unlike others, however, it provides an unbounded
range of distinct, discrete messages. Human language is acquired at a specific point
in development from within a limited range of possibilities, similar to the
acquisition of song in birds. Unlike the communicative signals of other species,
human language is under voluntary control, with its underlying neurobiology
concentrated in cortical structures, as opposed to the sub-cortical control
characteristic of those other species that have been studied in this regard.
Human language is structurally a discrete combinatorial system, in which elements
from a limited set combine in a recursive, hierarchical fashion to make an
unlimited number of potentially novel messages. The combinatorial structure of
language is governed by two quite independent systems: a small inventory of
individually meaningless sounds combine to make meaningful words, on the one
hand (PHONOLOGY), while these words are combined by a quite different system
to make phrases, clauses, and sentences (SYNTAX). These properties (discrete
combination, recursive hierarchical organization, and duality of patterning) are
not simply idiosyncratic ornaments that could in principle be omitted without
affecting the overall communicative capacity of the system. Rather, they are what
makes large vocabularies practical and unbounded free expression possible.
Contrast the unlimited range of potentially novel utterances which any (normal)
speaker of a language can produce, and another speaker of the same language
comprehend, with the strictly limited range of meaningful signals available to
other organisms. No other form of communication found in nature has these
properties. Human language, and especially its syntactic organization is quite
unique in the animal world.
Furthermore, efforts to teach systems with these essential properties to other
animals have not succeeded. Despite widespread claims to the contrary in the
popular literature, there is no evidence that any non-human animal is capable of

acquiring and using such a system. This should not be seen as particularly
surprising: if language is indeed embedded in human biology, there is no reason to
expect it to be accessible to organisms with a different biological endowment, any
more than humans are capable of acquiring, say, the echolocation capacities of
bats, a system which is equally grounded in the specific biology of those animals.

Human language is often considered as simply one more instantiation of the
general class of animal communication systems. Indeed, like others it appears to be
highly species-specific: although relevant experience is required to develop the
system of any particular language, the overall class of languages accessible to the
human learner is apparently highly constrained, and the process of language
learning more like genetically governed maturation than like learning in general.
The structural characteristics of human language are, however, quite different
from those of other communication systems, and it is the freedom of expression
subserved by those distinctive properties that gives language the role it has in
human life.

Stephen R. Anderson