T I N S - October 1985 453

Neuropathol. 60, 4 ~
17 Ohno, K., Pettigrew, K. and Rapoport, S.
(1978) Am. J. Physiol. 235, H299-H307
18 Fenstermacher, J. (1983) in Topics in
Pharmaceutical Sciences 1983 (Breimer, D.
and Speiser, P., eds), pp. 143-154, Elsevier,
Amersterdam
19 Pardridge, W. (1983) Physiol. Rev. 63,
1481-1535
20 Betz, A. L., Firth, J. and Goldstein, G.
(1980) Brain Res. 192, 17-28
21 Gjedde, A. and Christensen, O. (1984)
J. Cereb. Blood Flow Metab. 4, 241-249
22 Blasberg, R., Fenstermacher, J. and Patlak,
C. (1983) J. Cereb. Blood Flow Metab. 3,
8-32
23 Patlak, C., Blasberg, R. and Fenstermacher,
J. (1983) J. Cereb. Blood Flow Metab. 3, 1-7
24 Lund-Andersen, H. (1979) Physiol. Rev. 59,
305-352
25 Hawkins, Ra., Phelps, M., Huang, S. C.,
Wapenski, J., Grimm, P., Parker, R.,
Juillard, G. and Greenberg, P. (1984)
J. Cereb. Blood Flow Metab. 4, 505-517
26 Dunning, H. and Wolff, H. (1937) J. Comp.
Neurol. 67,433--450
27 Craigie, E. H. (1945) Biol. Rev. Camb.
Philos. Soc. 20, 133-146
28 Sakurada, O., Kennedy, C., Jehle, J.,
Brown, J., Carbin, G. and Sokoloff, L.
(1978) Am. J. Physiol. 234, H59-H66
29 Hawkins, Ri., Mans, A. and Biebuyck, J.
(1982) Am. J. Physiol. 242, E l - E l l
30 Bradbury, M. W. B. (1979) The Concept
of a Blood-Brain Barrier, Wiley, Chichester,
UK
31 Gross, P. M., Teasdale, G., Graham, D.,
Angerson, W. and Harper, A. M. (1982)
Am. J. Physiol. 243, H307-H317
32 Rapoport, S., Fredericks, W., Ohno, K. and
Pettigrew, K. (1980) Am. J. Physiol. 238,
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Joseph D. Fenstermacher is at the Departments
of Neurological Surgery and Physiology~Biophy-
sics, State University of New York at Stony
Brook, Stony Brook, N Y 11794--8122, USA.

Developmental aspects of infant speech

discrimination: the role of linguistic experience
Rebecca E. Eilers and D. Kimbrough Oiler
The h u m a n infant, once thought to be endowed with very limited auditory crimination paradigm was devised 4,5.
perceptual capabilities, has lately been credited with innate, complex speech The paradigm is based on an audio-
discrimination abilities. This article reviews and evaluates the evidence f o r innate logical assessment technique which
speech discrimination processing and examines the role o f specific language capitalizes on the infant's natural
experience in moderating innate abilities. proclivity to orient reliably to sound in
space by 4-6 months of age. In visually
Methods of evaluating infant speech becomes older and/or mobile. A more reinforced infant speech discrimina-
perception successful approach for assessing in- tion, the infant is rewarded (by the
The advent of infant speech percep- fants in this same age range capitalizes activation of an animated toy) for
tion research was made possible by the on the infant's interest in controlling turning toward a novel speech sound.
development of paradigms to assess his/her environment. The high ampli- The infant hears a repeating sound
the responses of non-verbal infants to tude sucking paradigm 2,3 allows the sequence such as 'ba,ba,ba' and is
speech and speech-like stimuli. Since infant to control the presentation rate taught to turn in anticipation of
the early 1970s three procedures, heart of speech stimuli by increasing his/her viewing the toy when the sound
rate, high amplitude sucking, and sucking rate. Sucking rate and ampli- changes, e.g., to 'pa,pa,pa'. The infant
visually reinforced infant speech dis- tude are monitored via a pressure is typically presented with 30 or more
crimination, have accounted for the transducer in a non-nutritive nipple. trials, during which the sound does
bulk of data. Once the infant learns that sucking (experimental) or does not (control)
The heart rate method of assess- results in the presentation of speech change. The activation of the toy when
ment of infant speech abilities is based stimuli, high-amplitude sucking rate the infant orients to the sound source
upon the cardiac orientating response 1 typically increases over a baseline during experimental trials serves to
and is applicable to infants between period. The increase.is followed sev- maintain the orienting behavior. The
birth and about six months of age. eral minutes later by habituation index of discrimination is the prop-
During this procedure, infants are (indicated by a sucking decrement); it ortion of turns to experimental v.
habituated to a repeatedly presented is at this point that a new stimulus is control trials.
speech syllable or word. After a introduced. Discrimination is inferred
number of presentations, the syllable if, after habituation, the infant's sucking Results showing infant speech
no longer elicits a cardiac deceleration rate increases relative to sucking rates discrimination abilities
(an indicator of infant attention), and of control infants when the new Since the advent of appropriate
the speech syllable is changed. Re- stimulus is introduced. assessment methods, it has been poss-
newed cardiac deceleration is taken as Although high amplitude sucking ible to show that infants can discrimi-
evidence of discrimination of the old has proven more robust than the heart nate among a number of speech con-
from the new speech sound. rate paradigm, its use is still limited to trasts differing across a number of
Although the heart rate paradigm infants in the first 4-6 months of important speech parameters. For
represented a methodological break- life. Consequently, neither high ampli- instance, infants can demonstrate dis-
through, practical limitations make its tude sucking nor heart rate provide a crimination of stop consonant syllables
use difficult. In particular, artifacts basis for particularly useful methods of differing in place of articulation (such
associated with gross motor activity assessing the effects of linguistic en- as 'ha' v. 'ga' or 'da' v. 'ga') in all three
often result in un-interpretable data, vironment over the prelinguistic paradigms 1,6-9. In these place-of-arti-
and, consequently, large numbers of period (birth to 12-18 months). In culation contrasts, the discriminative
infants must be evaluated and a great response to the need to study develop- cue occurs during a 25-40 ms period of
deal of data discarded. The problem mental aspects of perception, the acoustic frequency transition (format
becomes more serious as the infant visually reinforced infant speech dis- transition). Many fricative syllables
~) 1985.ElsevierSciencePublishersB.V., Amsterdam 0378- 5912/85/$02.00
454
(containing contrasts in high frequency
aperiodic sustained noise) such as 'sa'
v. 'va' and 'sa' v. 'sha' have also proven
discriminable1°. Similarly, discrimina-
tion of vowels 'oo' (as in 'boot') v. 'ee'
(as in 'beet') and 'uh' (as in 'but') v.
'ih' (as in 'bit') has been demonstrat-
edl~ .12.
Even more surprising than infants'
abilities to discriminate syllables is the
finding that infants perceive certain
speech sounds 'categorically'. Linguis-
tic categorical perception, as opposed
to continuous perception, is said to
occur when elements from two linguis-
tically distinct categories (e.g. 'ba' v.
'pa') are discriminable, but elements
from within the same category (e.g.
two acoustically distinct 'ba's') are not.
For the syllable 'ba' the lips part as
laryngeal vibration begins and the
vocal tract quickly assumes the posi-
tion for the vowel 'ah'. The time
between the lip burst and the onset of
voicing (i.e. the vowel beginning), the
voice onset time (VOT), is normally
less than 25 ms. Adult English-speak-
ing perceivers have a tendency to label
any such syllable with a VOT of less
than 25 ms as 'ba'. Similarly, syllables
with VOTs greater than 25 ms will
tend to be labelled as 'pa'. It is
possible, then, to have syllables from
the same category differ to a greater
extent (e.g. 'ba'-I with 0 ms VOT v.
'ba'-2 with 25 ms VOT) than syllables
from different categories along the
continuum ('ba' with 25 ms VOT and
'pa' with 40 ms VOT). One of the
advantages of categorical perception is
that it allows the perceiver to ignore
extraneous within-category informa-
tion and allows the speaker a fair bit of
articulatory variability without loss of
clarity. Infants, like adults, have been
shown to discriminate the categories of
English 'ba' and 'pa', represented by
synthesized syllables differing by only
20 ms of VOT. Infants have not shown
discrimination of two exemplars within
the category 'ba', synthesized with a
20-30 ms difference 3.13.
More recent work indicates that the
discrimination of syllables such as 'ba'
and 'pa' may not have been based
solely on the voice-onset time-contin-
uum. If the vocal tract reaches its
vowel target before voicing begins, the
resulting syllable will not include a first
formant transition reflecting the
change in vocal tract shape. Conse-
quently, syllables with long voicing lag
('pa's') often do not have a first
formant transition, while those with
short lag ('ba's') do 14. Apparently,
infants make categorical judgments of
voiced and non-voiced English-like
stop consonants using the presence or
absence of the formant transition to a
greater extent than voice onset time
per sd s. Changes in the duration of the
first formant transition produce
changes in the location of the categori-
cal perception boundary for infants. If
the formant transition is present, a
longer VOT is necessary to produce
the perceptual shift from 'ba' to 'pa '16.
Such results suggest that infants' abili-
ties to perceive speech sounds are
context sensitive.
An additional indication of the
context sensitivity of infant speech
perception is that syllable position
influences discrimination of conson-
ants 17. Other context effects can be
seen in a shift in the categorical
boundary between the consonants 'b'
and 'w' when the duration of the
syllable carrying the 'b' or 'w' is
altered. In longer syllables, longer
formant transitions are necessary for
the percept of 'w' than in shorter
syllables 18. Adult perception has been
shown to have similar properties of
context sensitivity 19.
The infant's capability to perceive
speech sounds categorically suggests
an awareness of natural categories of
speech sounds. A modified visually
reinforced infant speech discrimina-
tion procedure has been used to
demonstrate that infants group syll-
ables into phonemic categories by
ignoring irrelevant intonation, speaker
gender, and identityz°. There has been
an attempt to provide evidence that
infants have knowledge not only of
phonemic categories of speech sounds
(e.g. the categories used by a langu-
age to differentiate meaning as in 'pin'
v. 'tin') but also of the underlying
articulatory or featural similarity
across related categories (e.g. 'm', 'n'
and 'ng' are all nasal consonants and
'b', 'd' and 'g' are all oral conson-
ants) 21. Infants were presented with
groups of syllables that were featurally
similar or featurally mixed. Results
indicated that infant performance was
significantly better if the target syll-
ables were featurally similar. How-
ever, confounds in the experimental
design detract from the demonstration
and leave open the question of
whether infants have knowledge of
featural similarity2z.
The Innateness ItYi~.hesis
Based on early findings that infants
possess sophisticated speech process-
TINS October 1985
ing capabilities, several investigators
concluded that these abilities must be
innate. The Innateness Hypothesis 3
includes the following ideas: (1) in-
fants are innately capable of discrimin-
ating speech contrasts of many of the
world's languages; (2) infants discrim-
inate speech sounds in an adult-like
fashion; and (3) infant speech discrim-
ination is linguistic rather than audi-
tory. One implication of the hypothe-
sis is that development of speech
discrimination abilities may be un-
necessary because infants arc assumed
to have advanced skills. Consequently,
the early theoretical interpretations of
infant speech discrimination results
tended to inhibit interest in possible
effects of experience on infant speech
discrimination.
Criticism of the Innateness Hypo-
thesis first appeared in the mid-t970s.
The critics contested z~ the idea that
categorical perception, as indicated in
previous work 3, necessarily indicated
linguistic (as opposed to auditory)
perception. They noted that a variety
of crucial issues had not been resolved
by the existing studies and questioned
whether infant perception had been
shown to be linguistic or adult-like.
Shortly thereafter considerable inter-
est in possible effects of linguistic
experience on speech discrimination
abilities of infants began to appear in
the literature.
Results on effects of linguistic
experience
The Innateness Hypothesis did not
take into account the empirical evi-
dence that not all contrasts evaluated
had been shown to be discriminable by
infants. For example, the Spanish 'ba'-
'pa' contrast (which is substantially
different from the English one) ap-
peared to be very difficult to discrimin-
ate for English-learning infants.
Several contrasts that spanned the
Spanish category boundaries were not
discriminated23. These data are impor-
tant in the context of the claim that
infants can discriminate all 'universal'
contrasts, since the Spanish-style con-
trast appears to be as 'universal' (i.e.
as common across languages) as the
English one.
Additional studies 2.,25 empirically
addressed the possibility that the
failure of English-learning babies to
discriminate the Spanish voicing con-
trast might be due to a lack of relevant
listening experience. Both studies
(conducted in foreign countries with
non-English-learningbabies) indicated
TINS-October 1985
that infants from language back-
grounds that include the Spanish-like
contrast are successful in discriminat-
ing it. The studies inspired the inter-
pretation that listening experience
modifies infant speech discrimination
abilities very early in life. The inter-
pretation was bolstered by further
studies of both Spanish- and English-
learning infants in a single laboratory 13
which found that both Spanish- and
English-learning babies could discrim-
inate the English VOT contrast, but
only Spanish-learning infants dis-
criminated the Spanish VOT contrast.
The results of the studies on the
Spanish and English VOT contrasts
have suggested a role of experience
because Spanish-learning babies seem
to do better in discriminating the
Spanish contrast than English-learning
babies do. But why do the Spanish-
learning babies succeed with the Eng-
lish contrast? It has been suggested
that such success may be attributed to
an inherent salience of the English
VOT contrast, a salience that ove.r-
rides possible experience effects 14.
Even non-English-speaking adults are
able to discriminate the English VOT
contrast25.
Additional studies suggest that, in
some cases, the effect of experience
may be to inhibit the categorical
discrimination of an irrelevant con-
trast. Babies appear to discriminate
the English 'r' and T contrast, but
Japanese adults (whose language does
not include the contrast) show con-
siderable difficulties in appropriately
discriminating the same elements26. In
fact, the existing data indicate that
Japanese adults can discriminate in-
dividual exemplars of syllables with 'r'
and T, but not categorically, i.e. they
have difficulty identifying the proper
boundary. Recent results suggest that
the categorical boundary can be
trained in Japanese adults with a few
systematic sessions27. Similarly, it has
been found that infants lose the ability
to discriminate a Salish* speech con-
trast by the end of the first year of life
if they are not exposed to Salish 28.
Again the results suggest that in some
cases the contrast that is not supported
by experience may be lost between
infancy and adulthood.
The cross-linguistic studies of infant
speech perception, then, suggest an
extensive and varied role for experi-
ence in perception of speech sounds:
(1) discrimination of some contrasts
*Salish is a specific north-west American Indian
language of the Moan language family.
455
improves with experience during the rather than purely auditory. The fact
first year of life; (2) discrimination of that the chinchilla appears to dis-
some contrasts is little affected by lack criminate the English VOT contrast
of experience; and (3) discrimination categorically29 (very much like infant
of some contrasts disappears with lack and adult humans) provides further
of relevant experience. It is not known support to the idea that categorical
why different contrasts appear to be and adult-like perception of speech
differently impacted by experience or sounds may occur in an auditory rather
lack of it. than linguistic processing mode.
A more conservative model of
Results indicating that infant speech infant speech perceptual abilities re-
perception may be auditory rather quires eclecticism. It appears that
than linguistic several factors influence the discrimin-
Additional studies have contested ability of speech sounds in early
the claim that categorical or adult-like infancy. An important factor that was
perception may indicate a phonetic or not treated in the early literature is
linguistic mode of processing of speech intrinsic contrast salience or difficulty.
signals in infants. The chinchilla has Several studies suggest a hierarchy of
been shown to discriminate the Eng- discrimination difficulty that can be
lish VOT continuum much as infant correlated with degree of physical
and adult humans do 29. The chinchilla difference between stimuli32. Thus, a
data suggested that discrimination is variety of psychophysical results can
categorical and that the category be summarized by Weber's lawt.
boundary shifts under the influence of More recently, the asssumption of the
factors (place of articulation) that Innateness Hypothesis that categorical
shifts the boundary for humans. perception yields only between-cate-
Studies of the perception of non- gory discrimination independent of
speech stimuli by infants have also extent of physical difference has been
called into question the role of poss- empirically refuted with practiced
ible phonetic or linguistic processing. adult listeners33 but has not yet been
For example, one investigation3° of rigorously tested with infants. How-
infant perception of a nonspeech ever, it appears likely that the extent
continuum simulated the key transi- of physical difference and other qual-
tional and durational properties of an itative factors that may determine
earlier speech study using the syllables salience of auditory contrasts plays a
'ba'-'wa'ts. The results of the study major role in infant speech discrimina-
indicate that infants discriminate the tion.
nonspeech stimuli in the same categor- Finally, the effects of linguistic
ical manner as the speech stimuli. In experience must be accounted for by
both cases, the boundaries shift (to any model of infant speech perception.
similar locations) under the influence A variety of studies have now indi-
of stimulus duration. The results cated that perceptual abilities differ
suggest that infants' sensitive ability to across different infant language com-
perceive speech contrasts and to adjust munities. Experience appears both to
categorical perception by context may enhance contrasts that are heard by
be a more general auditory ability and the infant and to inhibit, in some
not a specifically linguistic one. cases, the discrimination of contrasts
that are irrelevant. Whether the
Trends in interpretation of infant changes involve peripheral auditory
speech perception restructuring or more central atten-
The results on early effects of tional factors is not yet determined.
experience and additional results on Innate abilities to discriminate
non-human speech discrimination re- speech sounds are clearly sophisti-
quire modification of the Innateness cated, but it is not clear how pervasive
Hypothesis3. It is true that many and how significant those abilities are.
speech contrasts are discriminable in Thus far, the weight of evidence
infancy, but some contrasts appear to suggests that innate abilities may be
be very difficult to discriminate unless auditory rather than phonetic. Addi-
experience intervenest3,31. Whether tional studies in the coming decades
infants discriminate like adults appears
to depend on what contrast and what t"Weber's law is the relation between stimulus
difference and the perception of those differ-
group of adults one considers, English, ences such that in observing the difference
Spanish, Hindi, etc. Finally, it is between two stimulus magnitudes, what is
unproven that early infant discrimina- perceived is the ratio of the difference to the
tion of speech sounds is linguistic magnitudes compared.
456
will serve to address the theoretical
gaps concerning the role of experi-
ence, salience, and specific auditory v.
linguistic processing in infant speech
perception skills.
Selected references
1 Moffit, A. R. (1971) Child Dev. 42, 717-731
2 Siqueland, E. R. and DeLucia, C. A. (t969)
Science 165, 1144
3 Eimas, P. D., Siqueland, E. R., Jusczyk, P.
and Vigorito, J. (1971) Science 171,303-306
4 Eilers, R. E., Wilson, W. R. and Moore,
J. M. (1977) J. Speech Hear. Res. 20, 766-
780
5 Eilers, R. E. and Gavin, W. J. (1981) in
Language Behavior in Infancy and Early
Childhood (Stark, R., ed.), pp. 185-213,
Elsevier, New York
6 Leavitt, L., Brown, J., Morse, P. and
Graham, F. (1976) Dev. Psychol. 12, 514-
523
7 Morse, P. A. (1972)J. Exp. Ch. Psychol. 14,
477--492
8 Williams, L. and Bush, M. (1978) J. Acoust.
Soc. Am. 63, 1223-1225
9 Eilers, R. E., Bull, D. H., Oiler, D. K. and
Lewis, D.C. (1985) The At-Risk Infant:
Psycho/SociolMedical Aspects (Harel, S. and
Anastasiow, N. J., eds), pp. 333--339, Paul
Comparative Neurology of the
Optic Tectum
edited by Horacio Vanegas, P l e n u m
Press, 1984. $125.00 (xx + 850 pages)
I S B N 0 306 41236 5
This h a n d s o m e volume has b e e n long
awaited by comparative vision enthu-
siasts and it rewards their patience by
providing a c o m p e n d i u m of detailed
information concerning tectal histol-
ogy, input and output pathways, neur-
onal response properties and bio-
chemical differentiation with represen-
tative studies in all five vertebrate
classes. This catalog of information
will be particularly useful in the library
of any d e p a r t m e n t with a strong
interest in comparative neurobiology
or in the evolution of visually guided
behavior.
O n the other hand, the volume has
notable deficiencies from the view-
point of many functional neurobio-
logists who see as neglected two m a j o r
needs of our emerging field: (1) a
better perception of the main trends in
brain evolution (what are firm h o m o -
logics, what is newly evolved?) and (2)
an understanding of how components
of the visual system can produce overt
behavior, adapted to the needs of the
organism. Given the long incubation
H. Brookes, Baltimore
10 Eilers, R. E. and Minifie, F. D. (1975)
J. Speech Hear. Res. 18, 158-167
11 Trehub, S. E. (1973) Dev. Psychol. 9, 91-96
12 Swoboda, P. J., Morse, P. A. and Leavitt,
L. A. (1976) Child Dev. 47, 459--465
13 Eilers, R. E., Gavin, W. and Wilson, W. R.
(1979) Child Dev. 50, 14-18
14 Stevens, K. and Klatt, D. (1974) J. Acoust.
Soc. Am. 55,653-659
15 Morse, P. A., Eilers, R. E. and Gavin, W. J.
(1982) Child Dev. 53, 189-195
16 Miller, J. L. and Eimas, P. (1983) Cognition
13, 135--165
17 Eilers, R. E. (1977) J. Acoust. Soc. Am. 61,
1321-1336
18 Eimas, P. and Miller, J. (1980) Science 209,
1140-1141
19 Miller, J. and Liberman, A. (1979) Percep.
Psychophys. 25, 457--465
20 Holmberg, T. L., Morgan, K. A. and Ktthl,
P. A. (1977) J. Acoust. Soc. Am. 62 (Suppl.
1), S ~
21 Hillenbrand, J. (1983) J. Speech Hear. Res.
26, 268-282
22 Moroff, D. A.(1985) J. Speech Hear. Res.
28, 316
23 Butterfield, E. and Cairns, G. (1974) in
Language Perspectives: Acquisition, Retarda-
tion and Intervention (Sehiefelbusch, R. and
Lloyd, L., eds), University Park Press,
of this volume, a reviewer could have
been found who could manage a
comparative chapter highlighting the
similarities and differences a m o n g the
major groups, or indeed the interest-
ing variations within groups. The
reader is given a fine view of the trees
(dendritic a n d otherwise) but only a
vague sense of the neuronal forest.
Because most authors in this volume
are anatomists and/or physiologists, it
might seem unfair to ask t h e m to
expound o n the relationships between
their findings a n d behavior. Yet many
of the issues that are raised by these
writers are treated inadequately, given
the background of long-available and
well-known articles by others. As one
example, G r a e b e r discusses interest-
ing effects of CNS lesions in sharks,
but without addressing the compara-
tive and functional implications of his
findings that tectum removal produces
less impairment of simple pattern
discrimination learning than does re-
moval of the telencephalon. Compar-
able experiments showing that goldfish
suffer little from ablation of telen-
cephalon (e.g. Savage) while they are
much impaired by tectum ablation
(e.g. Yager and Sharma) are not
mentioned by Graeber. The more
detailed studies on distinctive visual
17NS October i985
Baltimore
24 Streeter, L. L. (1976) Nature (London) 259,
39-41
25 Lasky, R. E., SyrdaI-Lasky, A. and Klein,
R. E. (1975)Z Exp. Ch. Psych. 20, 215-225
26 Miyawaki, K., Strange, W., Verbrngge, R.,
Liberman, A., Jenkins, J. and Fijimura, O.
(1975) Percep. Psychophys. 18, 331-340
27 Strange, W. and Dittman, S. (1984) Percep.
Psychophys. 36, 131-145
28 Werker, J. F. and Tees, R. C. (1984) Infant
Beh. and Devel. 7, 49~3
29 Kuhl, P. K. and Miller, 1. D. (1975) Science
190, 69-72
30 Jusezyk, P. W., Pisoni, D. B., Reed, M. A.,
Fernald, A. and Myers, M. (1983) Science
222, 175--177
31 Aslin, R. N., Pisoni, D, B., Hennessy, B. L.
and Perey, A. J. (1981) Child Dev. 52, 1135-
1145
32 Bull, D,, Eilers, R. E. and Oiler, D. K.
(1985) J. Acouat. Soc. Am. 77, 289-295
33 Carney, A. E., Widin, G. P. and Viemeister,
N. F. (1977)J. Acoust. Soc. Am. 62,961-970
Rebecca E. Eilers and D. Kimbrough Oiler are at
the Mailman Center for Child Development,
Department of Pediatrics, University of Miami
School of Medicine, P.O. Box 016820, Miami,
FL 33101, USA.
functions of the frog's tectum, pretec-
tum and thalamus (which provide a
background for discussion of parcella-
tion of functions in lower vertebrates)
are not mentioned here.
As a second example, Vanegas adds
to his elegant physiological review of
the teleost tectum a too brief sketch of
A k e r t ' s stimulation studies. Here he
uses a physiological parlance in re-
counting tpsiversive and contraversive
turns, rather than mentioning A k e r t ' s
hypothesis that the induced move-
ments resembled natural turns toward
food or flips away from threat. In this
context it would have b e e n appro-
priate to mention the complimentary
study of Springer et al. showing that
tectum ablation in goldfish abolishes
these two modes of orienting be-
havior. T h e one behavioral hypothesis
raised by Vanegas seems misplaced:
the ocular convergence produced by
A k e r t on stimulation of rostral tectum
of trouts was described as resembling
the 'lunge-and-snap' response trigger-
ed by sight of nearby frontal prey
(Akert) rather than 'food searching'
(Vanegas). The description by Vane-
gas of the failure o f tectumless fish to
catch food as a 'deficit in form
perception' is off the mark, especially
following Schneider's popular distinc-