The Development of Language in Bonobos and Chimpanzees

Pergamon
Language & Communication, Vol. 15, No. 2, pp. 121-148, 1995

Copyright 1995 Elsevier Science Ltd
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THE D E V E L O P M E N T OF L A N G U A G E SKILLS IN B O N O B O AND
CHIMPANZEE--I. COMPREHENSION
KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH
The species most closely related to our own are the bonobo (Pan paniscus) and the chimpanzee
(Pan troglodytes), who diverged from the hominid evolutionary path as little as five to eight
million years ago (Diamond, 1988; Miyamoto et aL, 1988). The two species of Pan, in turn,
shared a common ancestor two to three million years ago (Diamond, 1988).
Although chimpanzees and bonobos have demonstrated many complex cognitive skills (e.g.
Boysen and Berntson, 1990; Premack, 1976; Rumbaugh et al., 1993; Toth et al., 1993), their
capacity to acquire and use language has been a topic of controversy (Terrace et al., 1979;
Vauclair, 1990). Part of the difficulty in assessing language skills in apes arises from the fact
that they do not speak. Even if they possessed all of the cognitive requisites for language, the
structure of their vocal apparati would not permit articulate speech (Duchin, 1990; Lieberman,
1991). Thus, in order to examine language production in the great apes, investigators have
turned to sign (Fouts, 1973; Gardner and Gardner, 1969; Miles, 1983; Patterson, 1978) or to
artificial symbol systems (Premack, 1971; Rumbaugh, 1977; Savage-Rumbaugh, 1986;
Savage-Rumbaugh et al., 1986), each of which entail motor and/or vocabulary limitations for
the non-human subjects. Manual signs such as those used in American Sign Language (ASL),
for example, often require fine motor coordination that proves awkward for less dextrous ape
hands. Also, ape subjects using non-sign symbols are exposed to at most several hundred
symbols that they can produce (Premack, 1976; Savage-Rumbaugh et al., 1986) rather than the
thousands of words heard by most children as they mature. Evaluation of the apes' productive
skills is in large part, then, determined by modality limitations.
Comprehension may provide a better index of a species' competence for language than does
production. As far as we know, no peripheral anatomical limitations apply to apes' ability to
comprehend human speech. The ear structure and the sensory-perceptual auditory pathways of
the two genera appear to be very similar (Ankel-Simons, 1983); speech comprehension
differences between species, therefore, are likely to reflect CNS functioning. By evaluating
which of the receptive capacities underlying linguistic competence are well developed in
species closely related to our own, and which are not, we may be able to better understand how
humans came to be the adept linguistic creatures that we are.
Language comprehension in humans

Developmental psychologists are familiar with the premise that comprehension 'leads'
production (Goldin-Meadow et al., 1976; Huttenlocher, 1974). Comprehension of language
Correspondence relating to this paper should be addressed to Karen E. Brakke, Language Research Center, Georgia
State University, University Plaza, Atlanta, GA 30303-3083, U.S.A.
121
122
begins to be evident around 8-10 months of age (Bates, 1993), and a child's receptive
vocabulary early in the language-learning process is likely to be larger than its productive
vocabulary (Bates, 1993; Goldin-Meadow et al., 1976; Snyder et al., 1981).
Although language comprehension begins to appear early in life and does not inherently
require the fine motor abilities that speech and sign productive skills do, it is by no means a
simple or automatic phenomenon. The development of receptive language skills in our species
requires that several minimal conditions be met. Using spoken language as an example, the
individual must be able to distinguish different phonemes, then relate them to one another as
words. Meaning must be attached to the words, and finally the words must be systematically
related in order for the individual to derive the aggregate meaning of the utterance. Underlying
these linguistic skills are cognitive capacities such as the ability to integrate temporally or
spatially distinct pieces of information (Bates, 1993; Diamond, 1991), to mentally represent
absent entities (Piaget and Inhelder, 1969), and to establish part-whole relations (Langer,
1986). The context of the utterance and the communicative intent of the speaker are also
essential components of successful understanding of others' messages (Bruner, 1983; Lock,
1980).
In addition to cognitive requisites, appropriate environmental conditions must also be extant
for language-learning to occur. The infant cannot learn language unless it is exposed to it. It is
becoming increasingly evident that the development of language competence is heavily
supported by the people and culture surrounding the infant (Bruner, 1983; Lock, 1980). Use
of child-directed speech (Berko-Gleason, 1989; Ferguson, 1964; Snow, 1977) and joint
attention routines (Bakeman and Adamson, 1984; Bruner, 1975; Trevarthen and Hubley,
1978), for example, highlight the communicative process and engage the infant's attention.
Parents encourage children to participate in and respond appropriately to utterances during
daily activities long before they expect to hear similar utterances from the infants (Bruner,
1983; Goldin-Meadow et al., 1976).
Comprehension, then, arises in humans during normal day-to-day interactions between
infants and others. Children are not taught language by drilling repeated trials of words in
order to get a food reward. This must be considered when studying language competency in
other primates. In order to evaluate skills of related species at a level that permits comparison
to our own, we must provide rearing environments as similar as possible to those found in
human cultures. This includes providing linguistic and social input that is modeled after the
way human adults interact with infants around the world.
Language comprehension in bonobos

At the Language Research Center (LRC), two bonobos (Kanzi and Mulika), learned to use
and understand symbols in many different communicative capacities without explicit training
(Savage-Rumbaugh et al., 1990; Sevcik and Savage-Rumbaugh, 1994). Their instruction was
much less structured than that of other apes who were taught to use symbols in repeated-trial
or reward-based paradigms (Premack, 1971; Rumbaugh, 1977; Savage-Rumbaugh, 1986).
Language-learning by these bonobos involved observation of human models who used the
symbols as they interacted with the apes throughout the day. The young bonobos were asked
to attend to and participate in the daily activities of the laboratory (including play and
structured tasks), but were not required to produce symbols in order to receive food or other
rewards.
COMPREHENSION IN PAN
123
The two bonobos spontaneously developed receptive language skills and readily
comprehended several English words under blind test conditions (Savage-Rumbaugh et al.,
1993; Sevcik and Savage-Rumbaugh, 1994). The older of these bonobos has, in fact,
demonstrated the ability to understand several hundred novel complex spoken sentences
(Savage-Rumbaugh et al., 1993). Furthermore, the comprehension of symbols generalized to
symbol production in these subjects (Savage-Rumbaugh et al., 1990). In short, the two
bonobos were reared much like children and their symbol acquisition evidenced many of the
same characteristics as that of young humans.
Language-learning and routines
Central to the bonobos' learning of natural language was participation in interindividual
routines accompanied by adult language use. As outlined in Savage-Rumbaugh et al. (1993),
these routines are 'more or less regularly sequenced sets of interindividual interactions that
occur in a relatively similar manner on different occasions' (p. 25). These interactions are
embedded within a larger context. For example, within an 'evening' context when the lab is
quiet, there will occur routines involving milk preparation, quiet play, diaper changing, and
grooming until the infant falls asleep. Each of these routines, in turn, is made up of a sequence
of acts of components such as--for milk preparation--putting the lining in the bottle, opening
the milk formula, pouring the formula into the bottle, adding water, putting the nipple on, and
finally drinking the milk.
Within these routines, each participant has a role that from day to day stays relatively
consistent but over the course of time may shift, with the infant becoming more and more
active in initiating the components of the routine. Initially, however, the adult participant
directs the flow of the interaction and assigns a role to the infant that consists of paying
attention and indicating readiness to continue with the next component of the routine. Salient
components or transitions of the routine are behaviorally and linguistically (via speech and/or
lexigram) encoded by the adult. Such 'marking' of the components appears to be most
effective when it occurs prior to the actual event (Savage-Rumbaugh, 1991). Talking about
what we are going to do, before we do it, allows all participants to predict what will happen
next. Once able to predict upcoming events, the infant can begin to exert some control over
how those events unfold.
Quite often, components of a 'standard' routine will be added or modified. If, while
preparing milk, for example, it is discovered that the box of bottle liners is empty, a new box
will have to be located and opened. Linguistic marking becomes especially important at these
times when the routine is altered. By providing information that might otherwise be lacking,
the verbal information allows all of the individuals involved to coordinate their behavior so
that continuation of the routine may proceed smoothly (see Savage-Rumbaugh, 1991, for a full
discussion of the contributions of routines to language-learning in apes and children).
Species or rearing differences?

Prior to the study with bonobos at the LRC, two male chimpanzees, Sherman and Austin,
were taught referential use and comprehension of visuographic symbols ('lexigrams') via
explicit trial-based paradigms. Each communicative function--requesting, labeling, and
comprehension of others' utterances--was taught separately and required many repeated trials
for the chimpanzees to master. Productive skills did not generalize to comprehension without
explicit training (Savage-Rumbaugh, 1986), and comprehension of symbols, once learned, did
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KAREN E. BRAKKE and E. SUE S A V A G E - R U M B A U G H
not extend to speech (Savage-Rumbaugh et al., 1985b). Although Sherman and Austin
appeared to understand some verbal requests presented to them on a regular basis, when
compared on a formal single-word receptive language task ('Give me the x'), the two
chimpanzees demonstrated no comprehension of spoken English divorced of contextual
cues.
The data from Sherman and Austin on one hand and Kanzi and Mulika on the other
suggested a possible species difference in the ability to understand complex speech (SavageRumbaugh et al., 1985a). If confirmed, this difference would be a point of distinction between
the two species of Pan and might provide important clues to the evolution of human
language.
The comparison between the bonobo (Kanzi and Mulika) and chimpanzee (Sherman and
Austin) subjects was compromised by two differences in their rearing histories, however. The
chimpanzees were 1% to 2'~ years old when they started symbol training, while the bonobos
were 4-6 months old when they were first exposed to symbol use and English-speaking
caregivers. It may be that early exposure and attachment to humans is a critical component for
developing speech comprehension in chimpanzees.
There is some evidence that this may be the case. Fouts et al. (1976) reported appropriate
responses to at least 10 English words in a chimpanzee trained in ASL. Earlier studies with
chimpanzees Viki (Hayes, 1951) and Gua (Kellogg and Kellogg, 1933) suggested that some
speech comprehension may have developed in 'home-reared' subjects, but these reports were
based on data collected without contextual controls and so are inconclusive.
The second difference between the LRC studies involving the chimpanzees on the one hand
and the bonobos on the other was the language-learning environment experienced by the apes
after their inclusion in the project. The bonobos learned symbol use observationally, with an
emphasis on symbol comprehension. The chimpanzees, on the other hand, were first taught
symbolic request skills in a structured, trial-based paradigm. As Savage-Rumbaugh et al.
(1990) noted, it 'may be that an early emphasis on productive symbol t r a i n i n g . . , may, in
itself, interfere with acquisition processes that rely on observation' (p. 250).
In order to evaluate both productive and receptive language differences in the two species
of Pan, then, the next step was to raise a member of each species together from an early age,
providing them with the opportunity to learn language skills observationally and emphasizing
the comprehension of verbal and visual symbol use instead of requiring symbol production
from the outset.' This is the same paradigm used with the older bonobos, Kanzi and Mulika.
If under such conditions the chimpanzee fails to develop speech comprehension skills similar
to those that bonobos have exhibited, then this species difference may represent a key, abrupt
development in the phylogeny of linguistic competence. If at the other extreme, however, the
chimpanzee and bonobo profit equally from observational learning, this would testify to the
importance of early exposure to a linguistic culture in fostering comprehension skills even in
species other than our own.
This comparison has now been effected with a single bonobo and chimpanzee pair and the
results of the study relevant to comprehension are reported herein. Two general aspects of the
subjects' receptive language development provided the foci of the present study: their initial
entry into the system (i.e. how they first came to respond appropriately to utterances) and the
emergence of competence (i.e. the growth of their skills responding to single words and
complex utterances).
COMPREHENSION IN P A N
125
Method
Subjects
Panbanisha, the bonobo (Pan paniscus), was born at the Language Research Center (LRC)
on 17 November 1985. She was reared by her mother, Matata, until she was 7 weeks of age
when her care was transferred to human caregivers. Panpanzee, the chimpanzee (Pan
troglodytes), was born at the main station of the Yerkes Regional Primate Research Center on
31 December 1985 (the first chimpanzee born there after Panbanisha's birth) and transferred
to the LRC on 8 January 1986. From that day through the end of the study period (October
1989), Panbanisha and Panpanzee were nearly always together and in the company of at least
one human caregiver, except for the last months of the study when the subjects slept with older
bonobos at night. The subjects formed close attachment bonds with the caregivers who worked
with them consistently.
Rearing and communicative environment
Panbanisha and Panpanzee were reared together in an environment similar to that in which
many human children are raised, but one that took into account the activities and interests
to which young apes are prone. Eating, sleeping, bathing, changing diapers, traveling in
the woods and climbing trees, social interaction with other apes and humans, and quiet
'work sessions' (during which various assessments might be administered) filled the young
apes' day.
The different routines of the day, interspersed with occasional twists or surprises, lent
structure to the young apes' lives and also created an optimal environment for communication
about onging and upcoming events. Such an environment was considered crucial to the
emerging theory of non-human language acquisition that had been established a few years
earlier with the bonobos Kanzi and Mulika (Savage-Rumbaugh, 1991; Savage-Rumbaugh et
al., 1986). Panpanzee and Panbanisha were included in all aspects of laboratory life, and every
effort was made to convey information to them with lexigrams as well as with spoken English.
It was in this way, through observation of adult models and subsequent comprehension of
utterances, that the apes developed language skills.
Throughout the study, caregivers spoke to the subjects in English, much as they would
speak to human infants. No specific procedures were followed in speaking with the infants, but
speech could often be characterized as 'child-directed' (also known as 'motherese') (BerkoGleason, 1989), especially when the subjects were very small. Inflection, tone of voice, eye
contact, and gestures were all used as necessary to communicate the desired message during
daily interaction.
The principal difference between communication with these apes and communication with
human infants was that, whenever possible, visuographic symbols (called 'lexigrams') were
used to supplement or complement the English utterances. These symbols had been created
and assigned meanings during earlier study with other apes (Rumbaugh, 1977; SavageRumbaugh et aL, 1986). Each lexigram was glossed as the equivalent of an English word, such
as 'milk', 'dog' or 'good'. Approximately 256 lexigrams were available for use throughout the
study period. The symbols were arranged on faceplates throughout the lab but they
sometimes also appeared on individual photographs of the referents or were mounted
individually in appropriate places (for example, the 'bedroom' lexigram was taped to the
bedroom door).
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KAREN E. B R A K K E and E. SUE S A V A G E - R U M B A U G H
Data collection procedures

Daily data. Throughout the 4 years of the co-rearing project, the caregivers who
accompanied Panbanisha and Panpanzee throughout the day recorded virtually all instances of
symbol production and symbol and speech comprehension by each subject.: This was done in
order to provide an exhaustive record of all language-related behavior produced by the apes
that would be amenable to several types of analyses relevant to different aspects of language
acquisition. The Appendix provides examples of notes collected with the subjects during the
study period.
The daily data provided by the apes are similar to those of diary studies with human
children. Each entry was handwritten in a notebook as soon as possible after its occurrence
along with codes representing communicative function and behavioral concordance or
response, as well as contextual notes describing the context of the utterance or receptive
behavior. The entries were later entered onto a computer database, from which the receptive
data reported in this manuscript were extracted.
The receptive entries recorded in this way represent instances during daily interaction in
which a subject's comprehension of caregivers' utterances was evident. The exchanges
occurred within common daily routines and thus were often contextually appropriate.
Caregivers were, however, instructed to record only instances of comprehension that did not
appear determined or bound by extralinguistic cues including setting, gesture or glance. Thus,
for example, a request of 'get the soap' would not be recorded if the utterance was
accompanied by the adult's pointing or looking toward the soap.
Also, because most of the apes' caregivers had worked with other apes in similar conditions
and/or had worked consistently with Panbanisha and Panpanzee throughout the study period,
they were aware of responses that were made frequently and became routinized, and did not
record these behaviors as examples of comprehension. Information guiding the response had
to be judged as being gained from the words presented, or from textual rather than contextual
information. For example, an appropriate response to 'Use the towel' after taking a bath
would not be recorded if that is exactly what occurred at that same time every day anyway.
However, the same request and response used in a novel play situation, for example, would be
recorded.
Initially, from the start of the subjects' participation in early January through October 1986,
caregivers recorded observations of the infants' behaviors that they judged to be interesting or
significant. These notes were not systematic, but there were several references to the subjects'
communicative behavior that provided a good portrayal of their emerging communicative
competencies. Starting in late October 1986, daily receptive data were collected systematically
for both subjects through October 1989. The data set reported in this manuscript thus
represents, as completely as possible, the complete corpus of daily data collected during this
3-year period, from the first systematic recording of clear instances of comprehension when
the subjects were less than 1 year of age through the end of the study when the apes were
nearly 4 years old. A subset of these data was selected for the more detailed analysis of
sentence types reported in this manuscript; these were collected from January through October
1989. This period represents the end of the co-rearing project, thus the subjects' competencies
would be as highly developed as possible. The interval also coincides with the period of formal
testing of spoken English and lexigram comprehension reported below.
127
Utterances directed to the subjects consisted either of English only, or of English with
lexigram accompaniment. 3 All of the utterances entered in the data base were included in the
initial, general analysis for the two subjects. For the more detailed analysis of sentence types
(the 1989 data), only those entries that consisted of multiword utterances and were amenable
to syntactic analysis were included. These utterances could be categorized by semantic content
and were coded accordingly (e.g. Action-Object, Object-Action-Location, and so on).
Single-word test data. In addition to the daily record of utterance comprehension, a second
type of evaluation was conducted during the last year of the study period. This consisted of
controlled test situations in which subjects were asked to identify a spoken English word by
selecting among three lexigrams presented in unique arrays.
These assessments provided a decontextualized situation that could be used to literally test
or probe links between spoken word and graphic symbol that the subjects had formed in the
natural language-learning environment discussed earlier. The apes were required to generalize
knowledge gained in daily social interactions to very different circumstances.
These assessments can be likened to a multiple-choice vocabulary test. The response stimuli
consisted of the target plus two randomly selected alternatives. In both conditions, a standard
procedure was followed; the subjects had become familiar with this test situation in similar
assessments of spoken English and photographic stimuli the previous year. For each trial, E
gained the subject's attention, then presented the stimulus word either verbally or by playing
an audiocassette. E then opened a test booklet held between herself and the subject so that only
the subject could see the three alternatives placed within (see Fig. 1); E was blind to the
position of the target and each alternative. The subject was then asked to point to one of the
three lexigrams in the test booklet.
After the subject's response, E looked to see where S had pointed, judged whether the
response was correct, and provided appropriate verbal feedback (e.g. 'That's right!' or 'No,
that's not it. Sorry.') perhaps accompanied by a hug. If S pointed to multiple symbols, E asked
'Which one?' and asked for a clear, single response before looking and judging the response.
Continued multiple responses were counted as incorrect. Subjects were usually allowed to eat
or play quietly in between trials regardless of performance.
Videotaped records. In order to buttress the early notes of initial receptive episodes (i.e.
before notes were taken systematically) and confirm the course of the subjects' early
communicative development, videotapes from the period of May through December 1986
were reviewed. Thirty-seven tapes, typically recorded weekly or biweekly and each containing
1-2 hours of the subjects' activity, were examined for evidence of emerging speech
comprehension in the young apes.
Results
Early roots of comprehension
The first recorded instances of responses to language in each subject--albeit heavily
supported by context and tone of voice--involved responses to two types of regulatory
utterances at approximately 6-7 months of age. These utterances were 'no' and 'come here'.
Both were important in establishing behavioral limits for the young apes and were used
frequently throughout the day. With both demands, a clear response across situations was
expected by the adult, and if that response did not occur there were direct behavioral
consequences relevant to the situation at hand.
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KAREN E, BRAKKE and E. SUE SAVAGE-RUMBAUGH
Fig. 1. Panbanisha completes a vocabulary trial by (a) first listening to the spoken word presented via
headpohones, and (b) pointing to one of the available lexigrams after E lifts the screen. Panbanisha
keeps her finger on the lexigram until E can see and respond to her choice.
129
By August 1986 it was clear that caregivers expected Panbanisha and Panpanzee to respond
appropriately to both 'no' and 'come here', and that non-compliance was considered defiance.
Observations that the apes looked at the speaker and sometimes vocalized, moved away, or
engaged in display behavior while doing so, supported the caregivers' conclusions.
From this time on, Panbanisha and Panpanzee started emitting behaviors related to the
graphic symbols used around them. They often held their faces very close as the symbol was
being touched by an adult. Between 6 and 8 months of age, each infant went through a period
during which they frequently touched a symbol immediately after a caregiver had touched it.
Such behaviors suggested that the apes were starting to attend very closely to communicative
utterances, even if they did not yet demonstrate that they understood exactly what was being said.
Occasional responses to non-regulatory words began to appear when the subjects were 8-9
months of age. On several occasions during this period, Panbanisha emitted a vocal (e.g. foodbark) or behavioral response to 'milk' or 'go', sometimes responding to the use of the lexigram
and sometimes to the spoken English alone.
Panpanzee, slightly younger than the bonobo, continued to appear more dependent on
contextual support than Panbanisha during this time, sometimes changing her behavior in
response to new information regarding familiar routines. The daily notes from 6 August 1986
contain the following description: 'As we visit [the] dogs, Panzee seems to anticipate when I
touch OPEN, over and over, that we are going to let them out as we have the last two days.
Her hair [piloerects] and she immediately [runs] and [holds] tightly to Kelly'. (Panpanzee-known as 'Panzee' to her caregivers--was just getting to know the dogs at this time and was
apparently still afraid of them.)
Symbolic markers were regularly inserted into the ongoing stream of interaction with
Panbanisha and Panpanzee, signaling important transition points in the activity. The use of
symbols as markers to announce parts of routines had a predictive value and let the infant apes
anticipate upcoming events. This allowed the individual to make appropriate responses (such
as holding an adult tightly) and exert some control over what happened to her. Contextual
cues--the temporal and physical setting, the caregiver's posture, gestures and vocal
intonation--may have supported the infant's interpretation of the message to some degree, but
certainly did not determine it. The caregiver's topic easily could have been another activity or
object appropriate to the same setting.
During the last few months of 1986 changes in the behavior of both caregivers and the apes
themselves became apparent. Panbanisha and Panpanzee now engaged in noticeably improved
visual attention to the adult's behavior as the caregiver proceeded through the sequential steps
involved in talking about and carrying out routines. Joint attentional activities--including
looking at objects and even picture books--became more common than in earlier months, with
the infant apes often taking an active roll (by pointing to photos in a book or moving a
caregiver's hand toward a toy, for example).
For their part, caregivers became more demanding of the subjects, waiting until they had the
subjects' full attention before proceeding with communications or activities or asking the
subjects to fulfill parts of the routines themselves. If the subjects had difficulty doing so,
caregivers provided just enough guidance to achieve the goal at hand.
It was at this time that Panbanisha's and Panpanzee's receptive skills began to broaden. By
the end of January 1987, one or both infants had appropriately responded at least once to
130
'chase', 'bite', 'orangs' (orangutans), 'tickle', 'open', 'bye', 'bug', 'grape', 'Kelly', 't-room',
'rubberband', and 'water'. These responses all occurred within some context, naturally, but
caregivers judged none to be determined or bound by the context in which they occurred.
After this point, both subjects continued to respond appropriately to a greater variety of
utterances, with increasingly greater dissociation from contextual support. However, as will
become evident in subsequent sections of this manuscript, the bonobo Panbanisha consistently
demonstrated greater receptive understanding of symbols and speech than did the chimpanzee
Panpanzee, both within the context of daily activity and in controlled test situations. The
chimpanzee nevertheless did evidence comprehension of several words and sentences using
the different means of assessment.
Comprehension of complex utterances in daily interaction
During the 3-year period in which utterances were recorded, the bonobo Panbanisha
responded appropriately to 2616 of 2852 total recorded utterances (92%) while the chimpanzee
Panpanzee responded appropriately to 1596 (81%) of the 1825 utterances that were directed to
her and recorded. As Table 1 shows, the accuracy rate for each subject remained consistent
over the 3-year period. This suggests that at some level the caregivers were sensitive to the
changing individual competencies of the subjects and made requests of them accordingly.
Panbanisha received approximately twice as many sentences as Panpanzee during each of
the first 2 years of the study; this also suggests that the caregivers 'expected' more of the bonobo
than of the chimpanzee. Despite the increased demands, the bonobo consistently responded
with greater accuracy than did her chimpanzee counterpart ()~2 [1, 4655] = 121.52, p < 0.01).'
Table 1. Utterances presented to Panbanisha and Panpanzee 1986-1989
Number correct
Total number given
Percentage correct
Panbanisha
1986-1987
1988
1989
Total
671
1268
663
2602
719
1406
712
2852
93%
90%
93%
92%
Panpanzee
1986-1987
1988
298
593
365
718
82%
83%
578
1469
735
1818
79%
81%
1989
Total
Because the difference between the subjects' competencies was apparent, a subset of
sentences collected in 1989 was analyzed in further detail in order to locate possible sources
of dissimilarity. Sentences were coded with regard to the nature and number of 'key terms'
included. These consisted of action, agent, object, recipient, and location terms that the
subjects would need to understand in order to respond appropriately. A total of 713 utterances
were recorded for Panbanisha from January 1989 through October 1989. Of these, 513 met the
criteria for inclusion in the current analysis (i.e. entire utterance with multiple 'key terms'
recorded, specific action response required, speech and lexigram input only). Of the 738
utterances presented to Panpanzee during the same period 477 met the criteria and were
similarly analyzed. Most of the utterances that were dropped were recorded as single-term
131
utterances such as 'tickle' or 'bowl'. Because more of these were recorded for Panpanzee, her
data set was influenced slightly more than Panbanisha's by their exclusion from the analysis.
Duplicate (in both word content and modality) utterances were also dropped from the analysis.
The remaining 483 unique sentences presented to Panbanisha and the 381 unique sentences
presented to Panpanzee were then analyzed by input modality (English only vs English plus
lexigram) and sentence type (e.g. action-object, object-action-location). This was done to
determine whether we could identify some of the different components of language
comprehension that affected the performance of each subject.
Input modality. Of the 483 unique utterances presented to Panbanisha, 362 (75%) used
lexigram plus English input, while 121 were in spoken English only. Three hundred and
eighteen (83%) of Panpanzee's 381 sentences were in English and lexigram; only 62 were
presented in English only) Thus, the caregivers gave Panpanzee a lower proportion of
sentences in English only (Z2 [1,863] = 9.72, p < 0.01). In so doing they placed fewer demands
on her speech comprehension skills and provided relatively more lexical assistance with the
visuographic symbols.
Nevertheless, Panbanisha again appropriately responded to requests for action more often
than did Panpanzee, regardless of whether utterances were presented in English alone or
English with lexigram accompaniment. Results by input modality are summarized in Table 2.
Panbanisha responded appropriately on 90% of English only sentences and on 95% of English
plus lexigram sentences. Panpanzee responded appropriately on 78% of English only
sentences and 73% of English plus lexigram sentences. Thus, given that caregivers were
sensitive to the capabilities of the subjects and adjusted their input a priori, each subject
responded equally well to English sentences with or without lexigram supplements. The
bonobo, however, had only a few inappropriate responses while the chimpanzee had difficulty
with about one out of every four requests made of her.
Table 2. Comprehensionof different input modalities
Panbanisha
English only
English + lexigram
Number correct
Numberrecorded
Percentagecorrect
109
342
121
362
90%
94%
Panpanzee
English only
49
62
79%
English + lexigram
229
318
72%
Note: one utterancedirected to Panpanzee was presentedvia lexigramonly, with no speech
(she was partially correct; see text).
Receptive language competency reflects the interplay of several sensory, cognitive, and
motor systems. The differential rates of appropriate responses between subjects could arise
from several factors. For example, Panpanzee may simply not have understood as many words
as the bonobo, her responses may have been more contextually bound than Panbanisha's, she
may have had trouble with extracting important information from long complex strings of
words, or she may have become distracted more easily and thus would not attend closely to
the utterances directed to her.
Although her caregivers did consider Panpanzee the more distractible of the two subjects,
this type of attentional difference did not lend itself to direct analysis with this data base. Other
132
K A R E N E. B R A K K E and E. SUE S A V A G E - R U M B A U G H
factors, however, were examined by analyzing this group of sentences from 1989 as well as
more structured vocabulary tests administered throughout the study period.
Sentence type and complexity. In order to determine whether sentence type or complexity
affected P a n p a n z e e ' s responses, caregiver utterances were categorized according to
grammatical form. The sentence types along with correct, incorrect, and partially correct
responses for each subject are presented in Table 3. Percentage of sentences given in each
syntactic category is depicted in Fig. 2; the percentage correct response for each sentence type
is shown in Fig. 3.
Table 3. Sentencecomprehensionby grammaticalform January 1989-October 1989
Panbanisha
Sentence type*
Two-term
AO
OA
OL
AL
AR
Three-term
AOL
OAL
ALO
AOR
ARO
ARL
Other
Totals
Ct
PC
221
4
19
42
13
13
Panpanzee
NR
1
1
1
77
4
2
7
14
10
1
1
12
1
6
2__6
2
451
23
7
483 Total sentences
PC
NR
136
5
19
21
8
33
1
2
2
2
37
16
9
10
6
9
2
1
5
1
1
12
6
1
2
1
1__33
278
69
25
384 Total sentences
1
9
* 'Key term' codes: A = Action,O = Object, L = Location, R = Recipient.

t Responsecodes: C = Correct response, I = Incorrectresponse, PC = Partiallycorrect response,
NR = No response/ignore.
The distribution of sentence types was similar for the two subjects. Approximately half of
all sentences given were in Action-Object form, such as ' G I V E B O W L ' or 'Slap the ball'
(lexigrams are indicated by all-caps). Another 17% for each subject were A c t i o n - O b j e c t Location, for example, 'Take the orange to the group room.' Most other sentences comprised
two- or three-term combinations of action, object, location, and recipient. Sentence types that
were presented only rarely, such as ' O b j e c t - L o c a t i o n - A c t i o n ' or four-term utterances such as
' A c t i o n - R e c i p i e n t - O b j e c t - L o c a t i o n ' , are represented in the 'Other' category.
Panbanisha responded appropriately to at least 92% of each sentence type (or group of
related sentence types): her performance did not appear affected by the grammatical form or
length of the utterance. In fact, the low number of totally inappropriate responses made by the
bonobo precluded any log-linear analysis of the effect of sentence type because expected cell
values could not be raised to appropriate levels, even when different sentence types were
combined.
C O M P R E H E N S I O N IN PAN
133
260
m
b
200
180
R
e
o
o
100
d
e
60
d
0
AO
OL
AL
AR
AOL
OAL
ALO
ARO
AOR O t h e r
Utterance Type
Panbanleha
Panpanzee
Fig. 2. Total number of sentences that were presented to Panbanisha and Panpanzee that fit the criteria
for inclusion (see text). These sentences comprised the 1989 subset of utterances. The proportional
distribution of the different sentence types was very similar for the two subjects.
100%
P
e
80%
o
@
n
t
C
o
60%
40%
r
r
e
O
20%
t
0%
AO
OL
AL
AR
AOL
OAL
ALO
ARO
AOR Total
Utterance Type
Panbanlaha
Panpanzee
Fig. 3. Percentage of appropriate responses made to the different sentence types by the two subjects.
Panbanisha responded well to all grammatical categories, but Panpanzee had significantly more
difficulty with three-term sentences and Action-Object utterances than with the other two-term
categories.
134
Panpanzee, on the other hand, performed extremely well in response to some sentence types
but had many errors with others. Her percentage correct ranged from 57%
(Action-Object-Location) to 90% (Action-Recipient). In order to determine the significance
of differences between sentence types, Panpanzee' s data were submitted to log-linear analysis.
Because of the low number of sentences included in some categories, some sentence types
were grouped together in order to facilitate this analysis. By reducing the number of
categories, expected values for the remaining cells were raised to levels considered acceptable
for such analysis (Tabachnick and Fidell, 1989). The groupings were made based on types and
number of key terms included and are presented in Table 4. The 'other' category was dropped
from this analysis because it contained a disjoint group of sentences and its inclusion as a
category was meaningless.
Table 4. Panpanzee's performance by sentence type: log-linear analysis results
Sentence
types
Correct responses
AO + OA
OL
AL + AR
OAL + ALO
AOR + ARO
AOL + ARL
Other responses
AO + OA
OL
AL + AR
OAL + ALO
AOR + ARO
ARL + AOL
Observed
frequency
Expected
frequency
Adjusted
residual
141.00
19.00
29.00
19.00
17.00
39.00
134,55
16,00
24,00
20,36
19,64
49.45
1.522
1.482
2.050
-0.602
-1.184
-3.158
44.00
3.00
4.00
9.00
10.00
29.00
50.45
6,00
9.00
7,64
7,36
18,55
-1.522
-1.482
-2.050
0.602
1.184
3.158
Likelihood ratio chi-square = 16.97638, df = 5, p = 0.005.
Log-linear analysis supports the conclusion that the chimpanzee's performance varied as a
function of sentence type (G 2 [5, 381] = 16.98, p < 0.01). Review of adjusted residuals
indicates that she responded relatively well to Action-Location and Action-Recipient
utterances (e.g. 'GO SCRUBBY-PINE-NOOK' or 'WASH KAREN') and had particular
difficulty with the cluster of Action-Object-Location and Action-Recipient-Location
utterances (almost wholly attributable to AOL errors, e.g. 'CARRY the STRING to the
GROUP ROOM') No other adjusted residuals were significant, although the direction of the
residuals for correct responses to all two-term sentence types were positive while the residuals
for all three-term categories were negative.
Three of the four sentence types that Panpanzee had the most difficulty with (as indexed by
percentage correct) were three-term sentences. Even if the action term of such an utterance is
relatively simple, such as 'get' or 'give', there still remain at least two additional terms that
must be remembered and integrated as the action is carried out. Panpanzee's response patterns
indicate that she may have had difficulty with one or more of the steps required to remember
and/or coordinate the multiple terms in her responses. A review of the contextual notes
accompanying the entries reveals that Panpanzee often acted on the appropriate object but took
it to the wrong location or performed an inappropriate but familiar action on that object. For
COMPREHENSIONIN PAN
135
example, after retrieving a sweet potato from the refrigerator Panpanzee carried it to the sink
and washed it (as is often done) instead of putting it in the microwave oven as requested. This
pattern usually did not occur if Panpanzee already possessed the object in question or if it was
immediately available; she then completed the request appropriately. Her behavior suggested
difficulty in remembering or relating all of the components of the utterance. It seemed that if
the linguistic information became too complicated, Panpanzee fell back on her extant
knowledge of typical routines to complete what she thought was an appropriate action. In other
words, when challenged, contextual information overrode verbal or textual information for
Panpanzee.
These difficulties do not appear to be responsible for the chimpanzee's errors in the
Action-Object category. Most of these requests were syntactically relatively simple. Instead,
the high number and variety of such sentences presented indicates that some other factor such
as vocabulary size may have contributed to her errors. Data from controlled tests of singleword comprehension, reported below, are consistent with the results of the sentence analysis
and indicate that vocabulary size may indeed have contributed to her lower level of accuracy
in responding to utterances made during regular daily interaction.
Table 5. Single-word test results
Condition
Total
words
tested
Number of words meeting

75% criterion
Number of trials
correct
Panban
Panzee
Panban
Panzee
English to lexigram
217
179
(82%)
79
(36%)
603/733
(82%)
365/795
(46%)
Audiotaped English
to lexigram
76
71
(93%)
39
(51%)
224/247
(91%)
171/277
(62%)
English to lexigram
(retest)
8 (Pb)
23 (Pz)
7
(88%)
18
(78%)
22125
(88%)
61/78
(78%)
Tests of single-word comprehension

As she did with the utterances provided in natural settings, the bonobo Panbanisha
performed consistently well on the blind receptive vocabulary test. Panpanzee had more
difficulty although she performed well above chance levels and her vocabulary appeared to be
increasing in size throughout the study. The results of the two test conditions and the ages of
the subjects during each are presented in Table 5 and are summarized below.
English to lexigram. English to lexigram testing began in January 1989. A total of 217
symbols were tested with Panbanisha and Panpanzee. The bonobo Panbanisha was correct on
82% of the trials presented to her and met the 75% criterion (at least three out of four correct)
on 179 of the words; she was 100% correct (at least three of three correct) on 140 of these.
Panpanzee, the chimpanzee, selected the correct lexigram on only 46% of the trials, meeting
criterion on 79 of the symbols. She was 100% correct on 33 of these. Table 6 lists the words
in each subject's receptive vocabulary as determined by this means.
136
KAREN E. BRAKKEand E. SUE SAVAGE-RUMBAUGH

Table 6. English --~ lexigram receptive vocabularies, 1989
Panbanisha
Apple
Austin
Backpack
Bad
Ball
Balloon
Banana
Bedroom
Bite
Blackberries
Blanket
Blueberries
Book
Bowl
Bubble
Bug
Bunny
Butt
Butter
Cabinet
Camper Cabin
Can opener
Car
Carrot
Carry
Cereal
Chase
Cheese
Cherry
Chow
Clay
Clover
Coffee
Coke
Cold
Collar
Colony Room
Crisscross Comers
Dog
Egg
Fire
Flatrock
Go
Good
Goodbye
Gorilla
Grab
Grapes
Spin
Spoon
Staff Office
Stethoscope
Stick
Straw
Strawberries
String
Sue
Sue's Gate
Groom
Group Room
Gully Gushers
Hamburger
Hammer
Happy
Hat
Hide
Hilltop
Honeysuckle
Hot
Hotdog
Hug
Hurt
Ice
In
Jello
Jelly
Juice
Kanzi
Karen
Keepaway
Kelly
Key
Kiwi
Knife
Koolaid
Krista
Lana
Lemon
Lemonade
Lever
Light
Lighter
Liz
Log Cabin
Lookout Point
Mary Ann
Matata
Melon
Middle Test Room
Midway
Milk
Money
Monster
Mouth
Mushroom
Mushroom Trail
Surprise
Sweet potato
T-room
Throw
Tickle
Tomato
Toolroom
Toothbrush
Toothpaste
Towel
Nail
No
Noise
Noodles
Observation Room
Oil
Onion
Open
Orange
Orange drink
Orange juice
Orangutan
Outdoors
Paint
Panbanisha
Panzee
Paper
Peaches
Peanut
Peas
Perrier
Phone
Pillow
Pineapple
Plastic bag
Play
Popsicle
Potato
Privet berry
Quiet
Rain
Raisin
Refrigerator
River
Rubber band
Salt
Sandpile
Scare
Scrubby Pine Nook
Sherman
Shirt
Shop
Shot
Slap
Sleep
Snake
Soap
Sour cream
Treehouse
Turtle
TV
Umbrella
Vacuum
Velvet plant
Vitamin
Wash
Water
Wipes
137
Sue's Office
Sugar
Sugarcane
Toy
Trash
Tree
Yes
Yogurt
Apple
Austin
Bad
Balloon
Banana
Bite
Blackberries
Blueberries
Book
Bug
Bunny
Chase
Chow
Clover
Coffee
Ice
Jeannine
Kiwi
Kool-aid
Lana
Lighter
Raisin
Rock
M & M's
Melon
Middle Test Room
Midway
Mushroom
Soap
Sparkler
Stick
Straw
Strawberry
No
String
Observation Room
Oil
Onion
Sugar
Sugarcane
Sweet potato
Coke
Open
Tickle
Crisscross Comers
Dog
Fire
Foot
Goodbye
Groom
Hide
Honeysuckle
Hurt
Orange
Orange juice
Perrier
Pineapple
Play
Popsicle
Potato
Tomato
Toothbrush
Water
Yogurt
Panpanzee
Salt
Scare
Shirt
Snake
Privet berry
Quiet
This test provided the clearest demonstration that Panbanisha's receptive vocabulary was
more extensive than Panpanzee's. It is equally important, though, to note that the 3-year-old
chimpanzee exhibited comprehension of nearly 80 spoken words under controlled,
decontextualized test conditions, a performance that far exceeds that of other Pan troglodytes
similarly tested (Savage-Rumbaugh, 1986). Tests with Sherman and Austin, even as adults,
yielded only chance performance in response to any spoken words (Cerutti et aL, 1993).
Audiotaped English to lexigram. In order to further ensure that the performance of these
subjects was not dependent on features of the situation such as how the experimenter was
saying the target word, several words were subsequently evaluated using audiotaped English
presented over headphones. Again, Panbanisha's performance was consistent with earlier
tests. She selected the correct lexigram on 91% of the trials and met criterion on 71 of the 76
words. Panpanzee was correct on 61% of the trials, meeting criterion on 39 of the words in the
taped stimulus set.
Because Panpanzee's performance improved in the headphone condition relative to the
spoken English task, the words that each ape had failed in one condition and passed in the
other were retested without the headphones in September 1989 (i.e. this test was conducted
under the same conditions as the larger English to lexigram test set that began 9 months
earlier). This was done in order to determine whether Panpanzee's improvement could be
attributed to maturation or whether the headphone presentation actually gave Panpanzee some
advantage over the 'live-speech' condition by focusing her attention on the task more
effectively.
138
KAREN E. BRAKKE and E. SUE S A V A G E - R U M B A U G H
In this final subset, Panbanisha was tested on eight words (three or four trials per word) and
was correct on 22 of 25 trials. The only symbol on which she failed to meet criterion was
'food'. Panbanisha had also missed this word in the identical condition earlier but had passed
it in the audiotaped condition.
Panpanzee was retested on 23 words; she was correct on 61 of 78 trials and met criterion on
18 of the words. Four of the five words that the chimpanzee failed to identify at this time
('blanket', 'good', 'mushroom-trail', 'turtle'; the fifth was 'rain') were ones that she had
missed in the first 'live-speech' condition and had passed when listening to headphones. The
data suggest that both subjects were still experiencing some vocabulary growth during this
period, because in this final test they each performed better on most of the symbols they had
failed months earlier under identical conditions. However, a few words (such as 'rain') may
have been 'on the cusp' of comprehension at this time, and may have been subject to
fluctuating performance as evidenced by her 'mastery' on one occasion and 'failure' at a later
date.
Discussion
For the first time, members of two non-human species have been raised together from
infancy and provided with an environment that fosters development of non-speech language
competency. This allows direct comparisons to be made between members of these species
and suggests areas of inquiry that may prove fruitful for investigation of the language-learning
process in humans.
The data sets reported above, although collected independently using different techniques,
demonstrate remarkable convergence of results. Undoubtedly, the bonobo had the greater
receptive competency of the two subjects throughout this period of development. Her
performance was not affected by removing potential contextual support or by combining the
words she knew into complex sentences. Panbanisha's pattern of receptive development,
together with that reported for Mulika (Sevcik and Savage-Rumbaugh, 1994) and for Kanzi at
older ages (Savage-Rumbaugh et al., 1986), paints a portrait of bonobo language acquisition
that, in character, is not unlike that of children prior to 21/~years of age. Although the ape's
vocabulary size appears to increase more slowly than that of human children, both learn by
participating in routines with lexical markers used by an adult, and both undergo
decontextualization of symbol comprehension as they mature.
More surprising than the bonobo's high level of performance was the chimpanzee's
demonstration of speech comprehension. Although her level of accuracy remained lower than
Panbanisha's throughout the study period, Panpanzee nevertheless demonstrated speech
comprehension skills well beyond those of other chimpanzees similarly tested but differently
reared (Savage-Rumbaugh, 1986). These data provide strong testimony to the effects of
rearing on language-leaming. By immersing her in a language-rich culture early in infancy and
providing her with interaction and opportunities for observational learning----conditions very
similar to those experienced by virtually all humans during infancy--this chimpanzee was able
to develop rudimentary language comprehension skills.
Taken together, the data from these subjects indicate that many of the neural prerequisites
for basic language acquisition were in place millions of years before anatomical changes
(Lieberman, 1991), further neuronal growth and reorganization (Deacon, 1988), and cultural
growth facilitated their expression and elaboration. The differences found between the
139
bonobo, chimpanzee, and humans suggest, however, that the last six to eight million years
have been volatile ones for linguistic evolution. Even members of the two ape species which
have been distinct for less than three million years clearly show different potentials for
language learning.
But where do these differences lie? It does not appear that a simple dichotomy of speech
comprehension exists between the two species. Panpanzee clearly understood several words
without any contextual support. Rather, it seems that a far more complex array of systems is
conspiring to produce the pattern of results seen in this study. The components of these
systems, rooted in neurophysiological structure and process, may include, for example,
memory, attention, and ability to organize information.
Consider the role of attention, for example. In order to learn the meaning of a spoken word,
one must at some point attend to the sound being uttered as well as the referent of that sound.
If a lexigram is to be used as a visual representation of that word, then it must be accorded
some attention as well. Additionally, the presentation of all of these stimuli must allow the
individual to link them together as symbols and referent. The process requires the ability to
focus on the critical components of the situation (in this case, at minimum, the word, lexigram,
and corresponding object or event) and, if all are presented in temporal synchrony or
proximity, the ability to switch attention back and forth between components as well as the
speaker and context. If sources of relevant information are temporally distinct, such as when
a referent is absent when its corresponding symbol is used, the individual's long-term memory
also comes into play.
As an infant, Panpanzee was, compared with her bonobo and human peers, delayed by
several months in her ability to attend to more than one object at a time (Brakke and SavageRumbaugh, in prep.). This deficit may have carried over into the communicative domain,
making it more difficult for her than for Panbanisha at early ages to link symbols and their
referents. Although Panpanzee was better able to control her attention as she got older,
she still appeared easily distracted by extraneous events. The words for which the chimpanzee
demonstrated the strongest receptive skills were those for which she behaviorally
demonstrated the greatest interest. It may be that the introduction of these stimuli created a
motivational or arousal state that allowed Panpanzee to focus on what she wanted and thus
learn the critical relationships.
Another area in which critical species differences may be extant is the ability to process and
integrate sequential bits of information or information from multiple sources. These capacities
have been linked to frontal lobe functioning (Diamond, 1991; Goldman-Rakic, 1987). This
part of the neocortex is an area of the brain in which there are notable differences in volume
between human and apes (Passingham, 1982), although analogous comparisons between
bonobo and chimpanzee neocortices have not, to the authors' knowledge, been made.
The chimpanzee demonstrated difficulty in responding to utterances in which multiple
aspects, such as an action, an object, and a location, must literally be 'kept in mind'. It is
reasonable to expect such difficulties as a result of neural processing limitations with respect
to integration of different units of information (Diamond, 1991). Vocabulary acquisition, too,
requires integration of information provided via different sensory channels (see Bates, 1993).
Panpanzee may have had more difficulty than Panbanisha relating all the sources of
information necessary for successful word learning; thus her vocabulary at any given time was
not as extensive as the bonobo's.
140
K A R E N E. B R A K K E and E. SUE S A V A G E - R U M B A U G H
Although this discussion of factors affecting language comprehension has been presented in
the context of chimpanzee versus bonobo, it may well be relevant to the phylogeny of human
language. The same systems that appear to be limiting the chimpanzee--attention, memory,
processing and integration of information--may have allowed humans to proceed beyond the
limits currently observed in bonobos. For example, Panbanisha was certainly better able to
take advantage of her language-rich upbringing than was Panpanzee, but compared with a
3-year-old child the bonobo is limited in the complexity and variety of linguistic devices she
understands. To illustrate, at age 3 Panbanisha's known receptive vocabulary consisted of
approximately 200 words. Panbanisha may have understood some English words that were not
amenable to matching with lexigrams, but still the apparent size of her vocabulary did not
approach that of a normally developing human child of the same age. Similarly, although not
specifically tested during this time, the complexity of the utterances to which Panbanisha was
capable of responding was probably not as great as those a like-aged child could handle easily.
In fact, although Panbanisha' s older sibling Kanzi at 8 years understood a large set of requests
at approximately the same rate of accuracy as a 2-year-old human child, at the end of the study
several months later the child's accuracy had improved dramatically while Kanzi's remained
essentially stable (Savage-Rumbangh et al., 1993).
Conclusion
Although the P a n - H o m o relationship is not one of ancestor and descendant, comparative
results from bonobos and chimpanzees can provide clues about the possible order in which
different pieces of human language competence fell into place during our phylogeny.
It appears reasonable to assume that the skills that all three extant species have in common
were within the capacities of an ancestral form of anthropoid that lived around six or eight
million years ago. Thus, the ability to comprehend simple speech, understand referential
symbol use, and engage in intentionally communicative routines appear to have developed
prior to the emergence of language as we typically think of it. The expression of these skills
may have been dependent upon later appropriate cultural, anatomical and neural changes that
allowed recruitment, integration, and growth of all of the circuits implicated in language
competence (see Bates et al., 1979; Deacon, 1988; Ruse, 1988).
Among the components of language competence that seem to have come into place during
the last few million years, other than the speech production facility allowed by anatomical
modifications in the vocal tract (Lieberman, 1991), are the processes that allow an infant to
learn a very large number of words rapidly and to readily integrate multiple components of
various complex utterances while responding to them. Apes are not completely lacking in
these abilities, but the timing and extent of development do not appear to support language
development as fully as is commonly seen in humans. Growth and reorganization of the human
neocortex compared with the other species theoretically can account for many of the
phenomena associated with language and higher cognition (Deacon, 1988; Greenfield, 1991;
Jerison, 1973; Passingham, 1982).
Of course, Panbanisha and Panpanzee are only two individuals, and this study alone
contributes only one piece of the language puzzle. However, it is an important piece in that
Panbanisha's performance provides confirmatory evidence about the language-related
competencies of bonobos. Furthermore, Panpanzee's achievements demonstrate that
chimpanzees can learn to understand human speech and symbol use when immersed in a rich
communicative environment from early infancy. There are still, however, components of
COMPREHENSION 1N PAN
141
language-learning that appear to differentiate Panpanzee from her bonobo cousins, and future
investigation of these components may contribute to an understanding of the phylogeny of our
own language system.
Acknowledgements--This research was funded by National Institutes of Health grant NICHD-06016 which supports
the Language Research Center operated by Georgia State University. It was also supported by the College of Arts and
Sciences of Georgia State University. Support was also provided by grant RR-00165 from the National Center for
Research Resources to the Yerkes Regional Primate Research Center. The Yerkes Center is fully accredited by the
American Association of Laboratory Animal Care. Some of the data reported in this article were presented at the fifth
annual meeting of the Language Origins Society, Austin, Texas, 10-12 August 1989, or at the annual meeting of the
Eastern Psychological Association, Boston, Massachusetts, 3-5 April 1992. The authors wish to extend thanks to the
staff members of the Language Research Center for their excellent care of the subjects and invaluable assistance with
data collection.
NOTES
' The earlier bonobo subjects Kanzi and Mulika had, however, demonstrated spontaneous symbol production that
appeared to be facilitated by the opportunity to observe others using the symbols communicatively. Thus, symbol
production emerged in these subjects despite the fact that it was not 'taught'.
: A small number of utterances may have gone unrecorded owing to extenuating circumstances, such as when a pen
was lost while in the woods or when one person was single-handedly managing the behavior of and recording data for
two talkative chimps. Even in these situations, however, caregivers made 'mental notes' and recorded all utterances
they could recall as soon as possible.
Four utterances originally recorded during this period included gestures. Three of these were dropped from the
analysis. One utterance that included a gesture was retained. The gesture was not judged to contribute to the subject's
overall comprehension of the sentence; rather, it pointed out an unfamiliar object that the subject was not expected to
know.
' It must be acknowledged that placing increased demands on Panbanisha may have aided her performance---each
subject may have 'lived up' to caregivers' expectations. However, this interpretation would have to be tempered by
the knowledge that caregivers were highly motivated to help instill communicative competence in both Panbanisha
and Panpanzee, if for no other reason than that symbolic communication provides a powerful tool for management
and coordination of behavior. The caregivers were in constant communication with both subjects, and continually tried
to 'push' them along by challenging them with different communicative means, but people had to respond to the
subjects as individuals and adjust their communications accordingly. Otherwise, making a high number of requests
that Panpanzee, for example, did not understand would be disruptive because she would become 'frustrated' and noncompliant, interrupting the interactive harmony that successful communication can foster. It must also be remembered
that, based on previous work with chimpanzees, Panpanzee was not expected to develop nearly as much receptive
competency as she ended up doing; she far exceeded any expectations that may have been placed on her. Thus,
caregiver interaction with the apes was driven by the subjects themselves rather than by a pre-existing hypothesis. Of
course, further research with other chimpanzees is necessary to resolve this issue conclusively.
5 One lexigram-only utterance (no speech accompaniment) was presented to Panpanzee; this was 'GIVE POTATO
SHOT' in 1989. Panpanzee's response was partially correct; she retrieved a potato from another part of the room but
did not combine it with the syringe until after the action was demonstrated by the caregiver. Panpanzee then was very
interested in giving the potato a shot and did so several times.
REFERENCES
ANKEL-SIMONS, F. 1983 A Survey of Living Primates and their Anatomy. Macmillan, New York.
BAKEMAN, R. and ADAMSON, L. B. 1984 Coordinating attention to people and objects in mother-infant and
peer-infant interaction. Child Development 55, 1278-1289.
BATES, E. 1993 Comprehension and production in early language development (commentary on Savage-Rumbaugh
et al., 'Language comprehension in ape and child'). Monographs of the Society for Research in Child Development
58 (3--4, Serial No. 233).
BATES, E., BENIGNI, L., BRETHERTON, I., CAMAIONI, L. and VOLTERRA, V. 1979 The Emergence of
Symbols: Cognition and Communication in Infancy. Academic Press, New York.
BENEDICT, H. 1979 Early lexical development: comprehension and production. Journal of ChiM Language 6,
183-200.
142
BERKO-GLEASON, J. 1989 The Development of Language, 2nd edn. Merrill, Columbus.

BOYSEN, S. and BERNTSEN, G. 1990 The development of numerical skills in the chimpanzee (Pan troglodytes). In
Parker, S. T. and Gibson, K. R. (Eds), 'Language' andlntelligence in Monkeys andApes. Cambridge University Press,
Cambridge.
BRAKKE, K. E. and SAVAGE-RUMBAUGH, E. S. 1995 The growth of object manipulation in infant chimpanzee,
bonobo, and human. Manuscript in preparation.
BRUNER, J. S. 1975 The ontogenesis of speech acts. Journal of Child Language 6, 183-200.
BRUNER, J. S. 1983 Child's Talk: Learning to use Language. Norton, New York.
CERUTTI, D. T., SEVCIK, R. A., SAVAGE-RUMBAUGH, E. S. and RUMBAUGH, D. M. October 1993
Equivalence class structures in the existing languages of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus).
Poster presented at the meeting of Southeastern Association for Behavior Analysis, Chapel Hill, North Carolina.
CRAMER, D. L. 1977 Craniofacial morphology of Pan paniscus. In Szalay, F. L. and Luckett, W. P. (Eds),
Contributions to Primatology, Vol. 10, pp. 1-64. S. Karger, Basel.
DEACON, T. W. 1988 Human brain evolution: I. Evolution of language circuits. In Jerison, H. J. and Jerison, I. (Eds),
Intelligence and Evolutionary Biology, pp. 363-381. Springer, Berlin.
DIAMOND, A. 1991 Frontal lobe involvement in cognitive changes during the first year of life. In Gibson, K. R. and
Peterson, A. C. (Eds), Brain Maturation and Cognitive Development, pp. 127-180. Aldine de Greuter, New York.
DIAMOND, J. M. 1988 DNA-based phylogenies of the three chimpanzees. Nature 332, 685-686.
DUCHLIN, L. E. 1990 The evolution of articulate speech: comparative anatomy of the oral cavity in Pan and Homo.
Journal of Human Evolution 19, 687-697.
FERGUSON, C. A. 1964 Baby talk in six languages. American Anthropologist 66, 103-114.
FOUTS, R. S., CHOWN, B. and GOODIN, L. 1976 Transfer of signed responses in American Sign Language from
vocal English stimuli to physical object stimuli by a chimpanzee (Pan). Learning and Motivation 7, 458-475.
GARDNER, R. A. and GARDNER, B. T. 1969 Teaching sign language to a chimpanzee. Science 165, 664-672.
GOLDIN-MEADOW, S., SELIGMAN, M. E. P. and GELMAN, R. 1976 Language in the two-year-old. Cognition 4,
189-202.
GOLDMAN-RAKIC, P. S. 1987 Circuitry of primate prefrontal cortex and regulation of behavior by representational
memory. In Handbook of Physiology--the Nervous System, pp. 373-417. American Physiological Society.
GOLINKOFF, R. M., HIRSCH-PASEK, K., CAULEY, K. M. and GORDON, L. 1987 The eyes have it: lexical and
syntactic comprehension in a new paradigm. Journal of Child Language 14, 23-45.
GREENFIELD, P. M. 1991 Language, tools and brain: the ontogeny of hierarchically organized sequential behavior.
Behavioral and Brain Sciences 14, 531-551.
HAYES, C. 1951 The Ape in Our House. Harper, New York.
HUTTENLOCHER, J. 1974 The origins of language comprehension. In Solso, R. L. (Ed.), Theories of Cognitive
Psychology. Erlbaum, Hillsdale.
JERISON, H. J. 1973 Evolution of the Brain and Intelligence. Academic Press, New York.
KANO, T. 1990 The bonobos' peaceable kingdom. Natural History 99(10), 62-70.
KELLOGG, W. N. and KELLOGG, L. A. 1933 The Ape and the Child. McGraw-Hill, New York.
LANGER, J. 1986 The Origins of Logic: One to Two Years. Academic Press, Orlando.
LIEBERMAN, P. 1991 Uniquely Human. Harvard University Press, Cambridge.
LOCK, A. 1980 The Guided Reinvention of Language. Academic Press, London.
MILES, H. L. 1983 Apes and language: the search for communicative competence. In deLuce, J. and Wilder, H. T.
(Eds), Language in Primates: Perspectives and Implications, pp. 45~61. Springer, New York.
MIYAMOTO, M. M., KOOP, B. F., SLIGHTOM, J. L., GOODMAN, M. and TENNANT, M. R. 1988 Molecular
systematics of higher primates: genealogical relations and classification. Proceedings of the National Academy of
Sciences 85, 7627-7631.
PASSINGHAM, R. E. 1982 The Human Primate. W. H. Freeman, Oxford.
PATTERSON, F. G. 1978. The gestures of a gorilla: language acquisition in another pongid. Brain and Language 5,
72-97.
PIAGET, J. and INHELDER, B. 1969 The Psychology of the Child. Basic Books, New York.
PREMACK, D. 1971 Language in chimpanzee? Science 172, 808-822.
143
PREMACK, D. 1976 Intelligence in Ape and Man. Edbaum, Hillsdale.

RUMBAUGH, D. M. 1977 Language Learning by a Chimpanzee. Academic Press, New York.
RUMBAUGH, D. M., HOPKINS, W. D., WASHBURN, D. A. and SAVAGE-RUMBAUGH, E. S. 1993 Chimpanzee
competence for counting in a video-formatted task situation. In Roitblat, H. L., Herman, L. M. and Nachtigall, P. E.
(Eds), Language and Communication: Comparative Perspectives, pp. 329-345. Edbaum, Hillsdale.
RUSE, M. 1988 Intelligence and natural selection. In Jerison, H. J. and Jerison, I. (Eds), Intelligence and Evolutionary
Biology, pp. 13-33. Springer, Berlin.
SAVAGE-RUMBAUGH, E. S., 1984 Pan paniscus and Pan troglodytes: contrasts in preverbal communicative
competence. In Susman, R. L. (Ed.), The Pygmy Chimpanzee, pp. 395-413. Plenum, New York.
SAVAGE-RUMBAUGH, E. S. 1986 Ape Language: From Conditioned Response to Symbol. Columbia University
Press, New York.
SAVAGE-RUMBAUGH, E. S. 1991 Language learning in the bonobo: how and why they learn. In Krasnegor, N.,
Rumbaugh, D. M., Schiefelbusch, R. L. and Studdert-Kennedy, M. (Eds), Biological and Behavioral Determinants of
Language Development, pp. 209-233. Erlbanm, Hillsdale.
SAVAGE-RUMBAUGH, E. S., McDONALD, K., SEVCIK, R. A., HOPKINS, W. D. and RUBERT, E. 1986
Spontaneous symbol acquisition and communicative use by pygmy chimpanzees (Pan paniscus). Journal of
Experimental Psychology: General 115, 211-235.
SAVAGE-RUMBAUGH, E. S., MURPHY, J., SEVCIK, R. A., BRAKKE, K. E., WILLIAMS, S. L. and
RUMBAUGH, D. M. 1993 Language comprehension in ape and child. Monographs of the Society for Research in
ChiM Development 58 (3-4, Serial No. 233).
SAVAGE-RUMBAUGH, E. S., RUMBAUGH, D. M. and McDONALD, K. 1985a Language learning in two species
of apes. Neuroscience and Biobehavioral Reviews 9, 653-665.
SAVAGE-RUMBAUGH, E. S., SEVCIK, R. A., BRAKKE, K. E., RUMBAUGH, D. M. and GREENFIELD, P. M.
1990 Symbols: their communicative use, comprehension, and combination by bonobos (Pan paniscus). In RoveeCollier, C. and Lipsett, L. P. (Eds), Advances in Infancy Research, Vol. 6, pp. 221-278. Ablex, Norwood.
SAVAGE-RUMBAUGH, E. S., SEVCIK, R. A., RUMBAUGH, D. M. and RUBERT, E. 1985b The capacity of
animals to acquire language: do species differences have anything to say to us? Philosophical Transactions of the
Royal Society 3tl8, 177-185.
SAVAGE-RUMBAUGH, E. S. and WILKERSON, B. J. 1978 Socio-sexual behavior in Pan paniscus and Pan
troglodytes: a comparative study. Journal of Human Evolution 7, 327-344.

SEVCIK, R. A. and SAVAGE-RUMBAUGH, E. S. 1994 Language comprehension and use by great apes. Language
& Communication 14, 37-58.
SHEA, B. T. 1981 Relative growth of the limbs and trunk in the African apes. American Journal of Physical
Anthropology 56, 179-201.

SIBLEY, C. G. and AHLQUIST, J. E. 1987 DNA hybridization evidence of hominoid phylogeny: results from an
expanded data set. Journal of Molecular Evolution 26, 99-121.
SNOW, C. E. 1977 The development of conversation between mothers and babies. Journal of Child Language 4,
1-22.
SNYDER, L. S., BATES, E. and BRETHERTON, I. 1981 Content and context in early lexical development. Journal
of Child Language 8, 565-582.
TABACHNICK, B. G. and FIDELL, L. S. 1989 Using Multivariate Statistics. Harper-Collins, New York.
TERRACE, H. S., PETTITO, L. A., SANDERS, R. J. and BEVER, T. G. 1979 Can an ape create a sentence? Science
2116, 891-900.
TOTH, N., SCHICK, K., SAVAGE-RUMBAUGH, E. S., SEVCIK, R. A. and RUMBAUGH, D. M. 1993 Pan the
tool-maker: investigations into the tool-making and tool-using capabilities of a bonobo (Pan paniscus). Journal of
Archaeological Science 20, 81-91.
TREVARTHEN, C. and HUBLEY, P. 1978 Secondary intersubjectivity: confidence, confiding and acts of meaning
in the first year. In Lock, A. (Ed.), Action, Gesture, and Symbol, pp. 183-229. Academic Press, London.
ZIHLMAN, A. L. 1984 Body build and tissue composition in Pan paniscus and Pan troglodytes, with comparisons to
other hominoids. In Susman, R. L. (Ed.), The Pygmy Chimpanzee, pp. 179-200. Plenum, New York.
ZIHLMAN, A. L. and CRAMER, D. L. 1978 Skeletal differences between pygmy (Pan paniscus) and common (Pan
troglodytes) chimpanzees. Folia Primatologica 29, 86--94.
144
Appendix
Examples of daily notes

(Note: words written in all caps indicate that the lexigram for that word was used. Utterances in quotation marks are
ones that were made by the experimenter and directed to the subjects; these are the types of utterances included in the
analysis for this manuscript. The other entries, without quotes, are symbols touched by the apes. These will be
discussed in an upcoming report. The function code for each utterance is listed in parentheses after the utterance, and
the contextual notes appear as they were originally written, except that abbreviations have been written out and notes
corrected for tense agreement.)
Panzee
2 March 1989 PM
PEE
(statement)
While the chimps were sitting at the sink, listening to Sue talking at the keyboard, Panzee announced she
had to pee and then went to the potty.
GO
(imitation)
Sue was talking to the chimps verbally and at the keyboard about going to her house to visit the dogs. She
asked Panbanisha if she was interested in going--Panzee indicated she wanted to go by saying GO.
OBSERVATION-ROOM (request)
As we were about to go out the door to visit Sue's dogs, Panzee said OBSERVATION ROOM at the
keyboard. I wasn't sure if she was indicating she wanted to go there instead, or didn't understand about
going in Sue's car to visit her dogs at her house. However, the truck was gone, so we could not go up to
her house, so I suggested to Panzee that we could visit the observation room instead. She was interested in
doing this.
M & M (request)
In the observation room, Panzee said M & M and then led me to the door leading out of the room. Sue
mentioned to me that the chimps had seen Mary eating M & Ms through the window a few days ago, so
Panzee was probably referring to this. Mary walked by but told us she had no M & Ms with her today.
Later on Panzee and I went to the staff office and got some M & Ms there.
(behavioral note)
While in the observation room, Panbanisha walked to the end of the room and kicked the wall in a typical
'pygmy [chimp] display', as she looked towards the room where monsters supposedly live. Panzee walked
behind and imitated Panbanisha, kicking the wall in the exact same manner. I have never seen her kick
things in this way, and have only seen the pygmys kick like this.
PLAY
(imitation)
Panzee and Panbanisha started to play with each other in the observation room. I started to tell them
verbally and at the keyboard that if they wanted to play, they could do so in the group room. Panzee
interrupted me in mid-sentence and imitated PLAY to indicate she was interested in playing (see next
entry).
T-ROOM (incorrect request)
I finished my sentence about going to the group room to play, asking Panzee if she wanted to do this. She
replied T-Room. However, she led to the bedroom and wanted to play there instead.
PLAY STRING This/gesture (request)
After playing keep-away with some string for a while, I sat down to write some notes. Panzee indicated she
was still interested in playing with me and the string by saying PLAY STRING and touching the string.
This time I suggested playing chase with the string, and I chased after her with it.
'The STRING is HIDEing by the BLANKET.' (English + lexigram)
(inappropriate response)
I told Panzee I was going to hide the string, then went out of her sight to do it. I came back to the keyboard
and told her the above. She listened carefully, then took off running in the appropriate direction, but
eventually ended up in the toolroom. She searched the tool sites in there (see next entry).
'The STRING is HIDEing by the BLANKET in GROUP-ROOM.'
(English + lexigram) (appropriate response)
Panzee didn't seem to understand when I kept telling her about the string hiding by the blankets. However,
when I added the part about the group room, she led to the group room and to the shelf where the blankets
are. She quickly found the string underneath one of them.
STRING (request)
As Panzee and I were at the sink cooking, she said STRING to ask to go play with the string she had earlier.
I reminded her we were cooking now, and she could play with the string when we were done.
145
'Will you OPEN the BUTTER?' (English + lexigram) (appropriate response)

There was a container of butter, some bananas, a sweet potato, and a plastic bag of M & Ms out on the
counter when I asked Panzee to do the above. She responded appropriately.
PLAY
(request)
Panzee asked to play while we were cooking. I told her LATER.
GO
(request)
Panzee asked to go while we were cooking. I told her LATER.
'Will you CARRY the BUTTER to the REFRIGERATOR?' (English + lexigram) (partially appropriate response)
Panzee went to the refrigerator, but without the butter. I had to remind her with a pointing gesture that I
wanted her to take the butter with her.
'Will you CARRY the M & Ms to the MIDDLE-TEST-ROOM?' (English + lexigram) (partially appropriate response)
For one of the first times, Panzee understood that she was to take something with her to another location-generally, when I ask her this type of question, she takes only herself and goes to the location unless I
specifically point to what I want her to take. However, she picked up our bowl of food instead of taking the
M & Ms, but she did lead to the middle room.
PLAY CHASE (request)
Panzee had a few bites of food upon arriving in the middle room, then made this combination. I still had to
get some things prepared, so I told her she could play but I did not wish to chase.
CHASE (request)
Panzee played by herself for a while, then came back to the keyboard to ask me to chase. I told her no.
STRING (comment)
Panzee commented about her string, as she was playing with it (see next entry).
CHASE (request)
While we were working, Panzee asked me to chase her. I told her no, but volunteered to tickle, which she
settled for.
PLAY CHASE (request)
Panzee made this combination to ask me to chase her. I told her no.
PLAY
(request)
During a [vocabulary test] trial, Panzee asked to be excused to play. I told her LATER, after she finished
up.
PLAY
(request)
Panzee did one more additional trial and then asked to play again. I still wanted her to do another one, so I
told her LATER.
YES
(undetermined function)
Panzee activated YES and then tried unsuccessfully to activate a few other keys then looked to me for my
response (see next entry).
PLAY
(request)
I asked Panzee, 'Yes what?' and she replied PLAY. She had completed the last trial I had prepared for her,
so I told her she could go play.
ORANGUTAN (request)
Panzee said ORANGUTAN as we were in the middle of a trial. I told her we could discuss visiting the
orangs when she finished the trials I had for her.
TICKLE (request)
Panzee completed her last trial and asked to be tickled. I obliged.
DOG
(incorrect request)
I asked Panzee verbally and at the keyboard if she wanted to go to the colony room and visit the orangs
now. She behaviorally indicated she did and I asked her if she could answer at the keyboard. She replied
DOG, though. I told her we could not go.
PEE DOG (statement for PEE) / (incorrect request for DOG)
I repeated my question about going to visit the orangs and Panzee replied with this combination. She did
need to pee and went to the potty. I got the impression she really did want to see the orangs but was saying
DOG to refer to them, as she did end up leading to the colony room and to their cage.
ORANGUTAN (imitation)
I told Panzee again that we could not visit the dogs and told her we could visit the orangs in the colony
room. She imitated ORANGUTAN this time and led to their cage.
OPEN
(imitation)
At the orangs' cage door I asked Panzee verbally and at the keyboard if I should open it. She vocalized
affn-matively and said OPEN after me.
This/gesture M & M (request)
Coming back into the middle room after visiting the orangs, Panzee touched the bag of M & Ms and said
M & M at the keyboard to ask for some.
146
Panbanisha
19 August 1989 A M
That/gesture TV (request)
At the sink, Panbanisha takes my hand and leads me to the bedroom. Panbanisha then taps the TV screen,
then goes to the keyboard and touches the symbol. I agree to turn it on.
MILK
(request)
When I arrived in the morning Panbanisha got a milkface and asked for MILK. I tried to encourage
'YOGURT' instead but at the refrigerator we found several cans of milk and Panbanisha desperately
wanted some so I gave her a glass.
PAPER (request)
After Panbanisha drank the milk she touched PAPER and wanted my clipboard. I let her have a few sheets
of paper, the pen, and the clipboard on the counter.
PLASTIC-BAG (request)
Panbanisha finished with the paper and touched PLASTIC-BAG. I told her she could get one and she
retrieved some off the shelves.
PLASTIC-BAG GET (imitation)
When I queried if Panbanisha wanted to 'GET A PLASTIC-BAG' she first imitated PLASTIC-BAG and
when I repeated '?do you want to GET one?' (hoping for a yes or no answer) she imitated GET. I told her
'YES you can GET one')
'Put the "PAPER IN PLASTIC-BAG" ' (Panbanisha makes no response to utterance)
Panbanisha was playing with her plastic bag and her paper was nearby I made this suggestion and she sort
of tapped her paper but just kept playing with the bag. It seemed she did not quite know what I was
suggesting so she kept doing what she was doing.
COLONY-ROOM (request)
At the counter Panbanisha asked to go to the colony room. I told her we couldn't go in but we could
'LOOK'. We looked in and saw Bob cleaning the cages.
YOGURT This/gesture (imitation)
I asked Panbanisha if she wanted some yogurt and she imitated YOGURT and tapped the container but she
didn't want any.
TV
(request)
Panbanisha then touched TV and looked at me.
BEDROOM (imitation)
When I replied that there was 'a TV in BEDROOM' Panbanisha touched BEDROOM also.
TV
(response to query)
I then asked '?GO BEDROOM' and Panbanisha touched TV. I accepted this as an affirmative answer.
BEDROOM (request)
As I started running the bath water Panbanisha asked to go to the bedroom. I replied 'LATER. NOW
WASH.' After bath and physical we traveled to the bedroom,
CHASE (request)
When we entered bedroom we found Phil and Panzee there, Panzee was peeing, so Panbanisha touched
CHASE and went over to Phil. Phil agreed to chase her.
MIDDLE-ROOM (statement)
As Phil chased her Panbanisha touched MIDDLE-ROOM as she ran by and opened the middle room door
as an escape route.
YES
(response to query)
Panzee had asked to play chase with Panbanisha and Panbanisha spontaneously answered her YES. They
romped for quite a while.
CHASE (request)
Panbanisha eventually tired of playing with Panzee and said CHASE at the alltalk keyboard. When I asked
who she wanted to chase she walked over to Phil and touched him. He agreed.
STAFF-OFFICE (request)
After playing with Phil, Panbanisha asked to go to the staff office. When we did so she led there.
KEY
(response to query)
When Panbanisha touched STAFF-OFFICE, I asked '?GO STAFF-OFFICE.' Panbanisha replied KEY,
which I took as an affirmative answer. I used keys to open the door and we went to the staff office.
OPEN
(request)
When we got to the staff office Panbanisha asked to OPEN the door and we went to the staff office.
OPEN
147
(request)
When we got to the staff office Panbanisha asked to OPEN the lexan cover over the lab keyboard.
I suggested that we open a CABINET instead.
"CARRY COKE to the GROUP-ROOM via OUTDOORS' (appropriate response)
We had gotten some Coke out in the staff office and now I asked Panbanisha to carry the Coke back to the
group room via outdoors. We had come in through the colony room, but now Panbanisha retrieved the Coke
and carried it to the outside door.
GOOD
(comment)
As we worked on the chase task, Panzee and Phil walked through the other side of the group room.
Panbanisha wanted to leave the table but I asked her to keep working. She touched GOOD and I hugged
her and told her she was doing 'GOOD work'. We returned to the task.
BUBBLES (request)
At the work table Panbanisha touched BUBBLES and picked up the bubbles, looking at me. I said she could
have them.
'Let's blow some in your "MOUTH" (appropriate response)
Panbanisha had been playing with the bubbles when I made this suggestion. She opened her mouth and
waited for me to blow into it.
PEE
(request)
Panbanisha asked to go pee as we worked. I let her go to the potty.
'LOOK FOOD REFRIGERATOR' (English + lexigram) (appropriate response)
After we finished working on the computer I suggested we look for some more food in the refrigerator.
PHONE (request)
As we worked on other tasks while Panzee took a turn at the computer, Panbanisha touched PHONE.
I agreed and Panbanisha led to the phone.
COLONY-ROOM (request)
After talking briefly on the phone Panbanisha asked to go to the colony room but I replied we'd have to go
'LATER'.
TRAILER (request)
While working Panbanisha asked to go to the trailer. I told her we could go there when we went outdoors
later.
'GET HAT T-ROOM' (English + lexigram) (appropriate response)
We had just done a HAT lexigram matching trial and I asked if Panbanisha could get one from the t-room.
She went to the t-room alone and did so.
T-ROOM SCARE NOISE (request)
As we worked Panbanisha wanted to go back to the t-room (to get the monster mask, I believe) but I asked
her to do a series of trials first. When finished she touched NOISE and took my hand. We looked in the troom then noticed that the toilet in the bathroom was making noise.
'GIVE MONSTER APPLE' (English + lexigram) (appropriate response)
We had brought a monster mask to the table and I asked Panbanisha to give it some apple. Panbanisha
picked up a piece of apple from the table and put it in the mouth of the mask.
APPLE This/gesture (request)
After doing some more trials, Panbanisha asked for the apple scraps that were left on the table. I told her
YES.
CEREAL (comment)
As Panbanisha ate apple she tapped the cereal lexigram that she had named/matched earlier (with some
difficulty) and then touched the corresponding symbol on the alltalk keyboard spontaneously.
TRAILER (imitation)
After we finished working I offered a trip to the trailer. Panbanisha touched TRAILER, also wanting to go.
YES CARRY (request)
At the trailer door Panbanisha touched YES CARRY and looked to the door. I agreed to carry her inside
briefly.
TV
(request)
We saw [a visitor, 'K'] in the trailer but I asked Panbanisha to stay with me. Panbanisha asked K to come
watch TV with us and at the TV picked out a tape labeled 'MONSTER'. She watched K's reactions to the
tape closely.
TV This/gesture (request)
When I turned the TV off Panbanisha asked to watch more and pointed to a tape labeled DOG. We watched
briefly.
CARRY (request)
Panbanisha lost interest in TV and touched CARRY, then tried to sneak over to K. I restated my desire for
her to stay with me, however.
148
MILK
(imitation)
When I asked Panbanisha if she was ready to go get milk, she indicated she was.
GOODBYE (imitation)
I asked Panbanisha if she could 'TALK GOODBYE' to K, meaning vocally, but Panbanisha said
GOODBYE at the keyboard instead.

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Pergamon

Language & Communication, Vol. 15, No. 2, pp. 121-148, 1995

Language comprehension in humans

KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH

Language comprehension in bonobos

Species or rearing differences?

KAREN E. BRAKKE and E. SUE S A V A G E - R U M B A U G H

KAREN E. B R A K K E and E. SUE S A V A G E - R U M B A U G H

Data collection procedures

KAREN E, BRAKKE and E. SUE SAVAGE-RUMBAUGH

KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH

Total number given

* 'Key term' codes: A = Action,O = Object, L = Location, R = Recipient.

KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH

Likelihood ratio chi-square = 16.97638, df = 5, p = 0.005.

Table 5. Single-word test results

Number of words meeting

Tests of single-word comprehension

KAREN E. BRAKKEand E. SUE SAVAGE-RUMBAUGH

KAREN E. BRAKKE and E. SUE S A V A G E - R U M B A U G H

KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH

BERKO-GLEASON, J. 1989 The Development of Language, 2nd edn. Merrill, Columbus.

PREMACK, D. 1976 Intelligence in Ape and Man. Edbaum, Hillsdale.

troglodytes: a comparative study. Journal of Human Evolution 7, 327-344.

Anthropology 56, 179-201.

KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH

Examples of daily notes

'Will you OPEN the BUTTER?' (English + lexigram) (appropriate response)

KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH

KAREN E. BRAKKE and E. SUE SAVAGE-RUMBAUGH

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