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Science, New Series, Vol. 213, No. 4513. (Sep. 11, 1981), pp. 1275-1278.
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preferred the twigs that were lower in Littlefield, R. M. Potter, J . Mummul. 61, 734 7. R. H. Smith, U.S. For. Serv. Gen. Tech. Bull.
(1980). 1532 (1977).
monoterpene concentration (4). 5. K. B. Sturgeon, Evolution 33, 803 (1979); M. A. 8. For the December data, a stepwise regression
The extent to which a-pinene influ- Radwan and G. L. Crouch, J . Wildl. Munuge. analysis, with the MAXR option of the SAS
38, 32 (1974). statistical package [J. H. Goodnight, in SAS
ences the choice o f food by tassel-eared 6. A 2.0-g sample of cortical tissue was frozen User's Guide, J. T. Helwig and K. A.Council,
squirrels was investigated in another se- under liquid nitrogen, ground to a powder, Eds. (SAS Institute, Raleigh, N.C., 1979)], indi-
transferred with 50 ml of distilled water, and cated that a-pinene accounted for 29.5 percent
ries o f tests ia which subjects were al- steam-distilled with n-pentane for 1 hour. The of the variance ( P = ,013); the semipartials
pentane extract was dried over anhydrous (SR2)for 3-carene, P-pinene, and limonene were
lovved to choose between plain mash Na,SO,, and the pentane was removed (micro not significant ( P > .25). Similar results were
(made from ground sunflower seeds and Vigreaux column). When the volume of the obtained for the seasonally combined data with
extract was less than 0.1 ml, an internal standard a-pinene accounting for 34.5 percent of the
laboratory food) and mash laced with of 10.0 p1 of n-tetradecane in n-heptane (200 p1 variance ( P = ,006); again the semipartials for 3-
varying amounts o f a-pinene (9).Repeat- of C,,H,, in 10 ml of C7H,,) wa added, and the -
carede. B-oinene. and limonene were not sianifi-
volume was then increased to 0.1 ml with n- cant (P i ' . z s ) .
ed-measures analysis o f variance indicat- heptane. The concentrated extract was analyzed 9. The mash consisted of 50 percent ground sun-
ed that the a-pinene content affected by gas chromatography (F and M model 810), flower seeds and 50 percent mouse food
with dual-flame ionization detection and a 104-m (Charles River); 5 ml of safflower oil was added
food consumption FF(2, 14) = 3.72; glass capillary column coated with Carbowax to each 100 g of mash. The mash containing
20M). A 5.0-pl sample of the concentrated monoterpene was made up in concentrations of
P = .025, one-tailed test];as the amount extract with added internal standard was inject- 0.1, 0.3, and 0.5 ml of a-pinene per 100 g of
o f a-pinene increased from 0.1 to 0.3 to ed into a glass-lined injection port at a split ratio mash. On each testina dav. after 23 hours of
of approximately I00 : 1. The oven conditions food deprivation, e a c 6 squirrel was given two
0.5 ml per 100 g o f mash, the proportion were 5 minutes at 65"C, then increased 1" per food dishes; one contained 20 g of plain mash,
o f food consumed decreased from 0.45 to minute to 16S°C, and isothermal thereafter. He- the other 20 g of mash laced with a-pinene. Once
lium make-up gas was supplied at 30 mllmin at a squirrel began to feed from either dish, it was
0.41 to 0.24, respectively. 'The drop in the exit end of the glass capillary column. The given 20 minutes to eat; then both dishes were
co~~sumption at 0.5 ml per 100 g was data were recorded on a Varian CDS-I I I chro- removed and the amount of each type of food
matography data system, which gave the area eaten was recorded. Each squirrel received five
significant (Scheffe test; P < .05, one- percent composition for the various components trials at each of the three concentration levels.
and the area of each component relative to 10 The monoterpene was obtained from Glidden,
tailed test) and showed that higher con- percent of the area of the internal standard. Organic Chemicals Division, SCM Corporation,
cer~trations o f a-pinene caused tassel- Presentation of data in this manner permitted Jacksonville, Fla.
the evaluation of amounts of each component in We thank J. R. McKnight, J. C. Pederson, R. M.
eased squirrels to reduce their intake o f a the samples relative to the internal standard a s Potter, and K. Wuensch for their help with the
preferred foodstuff. well as the determination of the (relative) total study and A. J . Cooper, S. D. Berry, A. J.
amounts of all monoterpenes in each sample. Pantle, and J. C. Jahnke for reading the manu-
Thus the amount o f a-pinene is impor- Kovats' indices were determined on the glass scriot. Suooorted bv NSF - grants BNS 76-05069
tant in choosing which ponderosa pine capillary Carbowax 20M column at 70°C for the and'^^^ '76.80423.'
monoterpenes, and mass spectra were obtained Present address: Cooperative Institute for Re-
trees are to be used as food sources by from the total essential oil with a Carbowax 20M search in Environmental Sciences and Depart-
glass capillary column in a quadrupole gas chro- ment of Environmental, Population, and Organ-
tassel-eared squirrels. Other factors that matography-mass spectrometer (Finnigan mod- ismic Biology, University of Colorado, Boulder
may affect twig preferences are nutri- el 1015C) with a model 6000 data system [H. 80302.
Maarse and R. E. Kepner, J. Agric. Food
tional quality o f the cortical tissue and Chem. 18, 1095 (1970)l. 23 October 1980
possible synergisms between nutrients
and combinations o f secondary com-
pounds (mono- and sesquiterpenes).
Contextual factors such as the visual
appearance o f the trees, presence o f con- Spatial Knowledge and Geometric Representation in a
specifics, and pheromonal cues may also Child Blind from Birth
act as determinants in feeding tree selec-
tion i f the chemistry o f the tree is accept- Abstract. A series of experiments demonstrated that a congenitally blind 2'/r-year-
able. old child-as well as sighted but blindfolded children and adults-can determine the
R. C. FARENTINOS* appropriate path between two objects aJier traveling t o each of those objects from a
Department of Zoology, Ohio third object. This task requires that the child detect the distances and the angular
State University, Co1umbu.s 43210 relationship qf the jamiliar paths and that she derive therefrom the angle of the new
P. J. CAPRETTA path. Our research indicates that the locomotion ofthe young blind child is guided by
Department of Psychology, Miami knowledge. of the Euc.lidean properties of a spatial layout and by principles j))r
University, Oxjbrd, Ohio 45056 muking inferences based on those properties.
R. E . KEPNER
Department of Chemistry, W e have had the opportunity to study Sighted adults and 3-year-old children,
University of Calij))rnia, Davis 95616 the spatially guided locomotion o f one all blindfolded, performed with similar
V. M . LITTLEFIELD21/2-year-old blind child in several ex- accuracy. These observations demon-
Department of Psychology, perimental settings. After the child had strate that the locomotion o f children,
Miami University been taken along several paths connect- with or without visual experience, is
References and Notes
ing four objects in a small room, she was guided by metric knowledge o f space.
able to move directly between the ob- This knowledge makes possible the deri-
1. J. 0. Keith, Ecology 46, 150 (1965); D. R.
Patton, U.S. For. Srrv. Res. Note R M 272 jects along paths she had never taken. vation o f further spatial information.
(1974). These observations were undertaken
2. D. R. Patton and W. Green, U.S. For. Serv.
Re.$. Note R M 169 (1970). to address a classical issue in psycholo-
3. Feeding trees were determined in the field by Fig. 1. Room layout gy, the development o f human knowl-
extensiveness of defoliation and number of
freshly clipped needle clusters; the nearest for spatial inference edge o f space. Descartes (1) suggested
neighbor of the same age and size a s the feeding experiment. The
tree that did not show cvidence of squirrel- room measured 2.44 that the geometric principles underlying
caused defoliation was designated a s a nonfeed- m by 3.05 m. Dashed spatial knowledge are innate and accessi-
ing tree. In choice experiments and monoter-
pene analysis, twigs from a specific feeding tree lines, trained routes; ble to any perceptual mode. He offered
were compared with twigs from its counterpart solid lines, test the example o f a blind man exploring
nonfeeding tree. routes. Landmarks:
4. P. J. Capretta and R. C. Farentinos, in Prefer-
M, mother; P, pil- objects with a stick. For the man to
ence Behuviour und Chemorrception, J. H. A. discover the shapes and arrangement o f
Kroeze. Ed. (Information Retrieval Limited, lows; T, table; B , bas-
London, 1979), pp. 205-218; . , V. M. ket. those objects, he must refer each tactual
SCIENCE, VOL. 213. I 1 SEPTEMBER 1981 0036-807518110911- 1275$01.0010 Copyright 0 1981 AAAS 1275
impression to a unitary spatial frame- they apply these inference rules to the tal fibroplasia. She is totally blind (5).
work, structured by the principles of sensory information that the environ- After spending the first 5 months in the
Euclidean geometry. Descartes proposed ment provides. Second, Gibson (4) pro- hospital, primarily in an isolette, Kelli
that any perceiver, exploring the world posed that no geometric principles what- was discharged wlth a developmental
through any mode, faces the blind man's ever need structure one's sensory im- status of newborn.
problem, and must draw on tacit knowl- pressions. 'The spatial properties of ob- When Kelli was 32 months old, she
edge of geometry to solve it. Descartes jects are specified by higher-order was brought into an unfam~liarlabora-
further reasoned that geometric princi- relationships in spatially and temporally tory playroom, 2.44 m by 3.05 m. The
ples must be innate: since those princi- extended arrays of stimulation. Space room contained four landmarks in a dia-
ples structure any perceptual experi- perception depends on mechanisms that mond shaped array: her mother seated
ence, they cannot themselves be ac- detect these relationships. on a chair (M), a stack of pillows (P), a
quired through experience. These argu- We have attempted to address these basket of toys (B), and a table (T) (Fig.
ments form part of the core of the issues by extending Descartes's exam- I). In experiment 1, we placed her at M
rationalist tradition in psychology (2). ple. We focused on the attempts of a and walked her from there to P and back,
Descartes's analysis can be contrasted young blind child to discover the spatial twice; from M to T and back, twice; and
with two major subsequent develop- layout of objects in a room from limited, from M to B and back, twice. Each time,
ments. First, Helmholtz (-?), extending temporally extended encounters with Kelli felt the object while facing it, and
the arguments of British empiricism, re- those objects and with paths between she might have thus determined the ori-
jected Descartes's claim of innateness. them. We investigated whether this child entation of the object relative to the
Helmholtz suggested that the geometric could make spatial deductions that rely trained path. We then induced her to find
principles underlying space perception on information about Euclidean angles the routes between P, T, and B on her
can be acquired. Perceivers need be en- and distances. own, by giving her such simple com-
dowed only with a set of general, non- Our principal subject was Kelli, a child mands as "Go to the toybasket" or "Go
spatial inference rules. They will deduce born 3 months prematurely and blinded to the pillows." Kelli was followed as
the principles of Euclidean geometry as shortly after birth as a result of retrolen- she moved until she reached the goal or
I I SEPTEMBER 1981
inally published in 1948). These geometries can- man for extensive overall conceptual guidance; rial disease [K~llner'srule ( 4 ) ] , indicat-
not support the inferences we have studied, U. Neisser for comments on a previous draft of
however, for they preserve no metric properties this paper; and K. Feldman for running sighted ing an undue vulnerability to retinal in-
IR. Courant and H. Robbins, What is Mathe- contfol sub'ects and transcribing the videotaped sult,
matics? (Oxford Univ. Press, New York, 1941)l. sess~ons.$his work was aided by a social and
8. We thank C. R. Gallistel, for focusing our atten- behavioral sciences research grant from the Na- In 1970, Laties and Liebman (5) re-
tion on formal geometric analysis of our experi- tional Foundation-March of Dimes and a Wil-
ment, for suggesting important controls and liam T. Carter Foundation grant, both to L. R. ported that the intravitreal injection of a
variations on the basic experimental paradigm, Gleitman. tissue-reactive fluorescent dye, Procion
and for suggesting a statistical analysis derived
from the avian navigation literature; L. R. Gleit- I5 December 1980; revised 22 May 1981 yellow, stained the outer segments of
cones, but not of rods, in the amphibian
retina. Using greater amounts of the dye,
we have obtained a striking result. In the
retina of the monkey, not only are all
Staining of Blue-Sensitive Cones of the Macaque cone outer segments stained with Pro-
Retina by a Fluorescent Dye cion yellow, but the entire soma of some
cones is completely stained by the dye,
Abstract. Intravitreal injection of aJluorescent dye, Procion yellow, results in the producing a Golgi-like silhouette. Such
complete and systematic staining of a cone population in the monkey retina. These cones are organized in a rather regular
cones form an approximately regular array whose separation varies with retinal array and have a characteristic retinal
eccentricity. They are absent in the very center of the fovea, and their density peaks distribution.
at I". The distribution of stained cones resembles that reported for blue-sensitive Procion yellow M4RAN (Polysci-
cones of other primates and, consistent with such an identijication, they are found ences), 5 to 7 percent in deionized water,
with less incidence in species having lower concentrations of blue cones. was injected (0.15 ml) intravitreally into
the eye of anesthetized rhesus and cyno-
The neural retina is a highly organized caque monkeys, the green and red cone molgus monkeys. Leakage of dye was
structure with a crystalline-like array of systems have similar electrophy siologi- reduced by the slow removal of the nee-
tightly packed cones and rods in a two- cal and psychophysical properties, dle. In some animals we also injected
dimensional matrix. Microspectrophoto- whereas those of the blue cone system Lucifer yellow (Polysciences) ( 6 ) simul-
metric work has provided evidence of are different (2). There are also differ- taneously with or subsequently to the
three cone types in the primate retina, ences between dysfunctions of these Procion dye in a weight ratio of 1 : 50 to
each having peak sensitivity at a differ- cone systems leading to color vision dis- 1:100 of Lucifer to Procion yellow. Ex-
ent part of the spectrum (1)-"blue-," orders. Congenital disorders of the green cept for one monkey, which was kept in
'6
green-," and "red-sensitive" cones. and red cone systems correspond to sex- the dark during and after the injection,
To our knowledge, morphological differ- linked inheritance, whereas those of the the animals were kept in a normal light
ences among these cone types have not blue cone system correspond to autoso- (200 trolands): dark cycle for 18 to 30
been reported in primates. Functionally, ma1 inheritance (3). In addition, blue hours (7). The animals were then killed
however, blue cones have unique prop- cones are often involved in acquired with an overdose of pentobarbital. The
erties. In humans and (Old World) ma- color vision disorders secondary to reti- eyes were fixed, often by arterial perfu-
Fig. 1. (A) Radial section of rhesus monkey retina (- 20' eccentricity) showing a cone completely stained by Procion yellow among other cones
unstained except for their outer segments. (B) Tangential section passing through the outer limiting membrane showing a regular array of stained
cones; unstained cones and rods appear as holes in the stained mesh of the outer limiting membrane.
1278 0036-807518110911-1278$01,0010 Copyright O 1981 AAAS SCIENCE, VOL. 213. I 1 SEPTEMBER 1981