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Cyclical Changes of Vaginal Cytology in the Cat

Article in The Canadian veterinary journal. La revue veterinaire canadienne · May 1979
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Cyclical Changes of Vaginal
Cytology in the Cat
JENNIFER N. M ILLS,
V.E. VALLI AND
J.H. LUMSDEN*
SUMMARY
Vaginal smears from seven cats were examined at
two-day intervals for 32 days in order to describe
the cyclical pattern of epithelial cells exfoliated
throughout the stages of the estrus cycle. Vaginal
epithelial cells were classified as parabasal, inter-
mediate and superficial (nucleate and anucleate)
cells, and their dimensions were measured for the
purpose of definition. The percentages of the
epithelial cell populations (i.e. Maturation Index)
from Wright's stained smears, were determined at
all stages of the estrus cycle. The Eosinophilic
Index was estimated on Papanicolaou stained
smears.
Smears of cats in estrus were populated almost
entirely with nucleate and anucleate superficial
epithelial cells. Proestrus was characterized by
intermediate epithelial cells with increasing eosino-
philia, and rare neutrophils. Metestrus was asso-
ciated with desquamation of intermediate and
parabasal epithelial cells, neutrophils and debris.
In the anestrus period, groups of intermediate cells
and some parabasal epithelial cells were exfoliated.
Two cats in the study did not cycle and exhibited
anestrus. Of the five cats cycling, eight estrus
periods were observed of two to five days duration.
The cycles were of 15 to 17 days interval in three
normal cats. Two cats did not show a second estrus
within 30 days, and were subsequently found to
have bacterial growth on the culture of vaginal
swabs, however the presence of an initial ovular
estrus cannot be ruled out. The rare presence of
erythrocytes was associated with vaginal bacterial
infections and discharge in two cats.
R SUM
Les variations cycliques de la cytologie vaginale,
chez la chatte
Cette etude portait sur sept chattes et consistait A
en examiner des frottis vaginaux, A intervalles de
deux jours et durant une periode de 32 jours, afin
d'etablir un profil d'exfoliation des cellules epi-
*Department of Pathology, Ontario Veterinary College,
Can. vet. J. 20: 95-101 (April 1979)
theliales au cours du cycle oestral. On classifia les
cellules de l'epithelium vaginal, selon leurs di-
mensions, en cellules parabasales et intermedi-
aires, ainsi qu'en cellules superficielles nucleees et
anuclees. Pour chacune des periodes du cycle
oestral, on determina le pourcentage des diverses
varietes de cellules, i.e. l'indice de maturation, A
l'aide de frottis colores selon la methode de Wright.
On determina par ailleurs l'indice eosinophilique, a
l'aide de frottis colores d'apres la methode de
Papanicolaou.
Les frottis vaginaux des chattes en oestrus
contenaient presqu'exclusivement des cellules su-
perficielles nuclees et anuclees. Le proestrus se
caracterisait par la presence de cellules inter-
mediaires, de quelques neutrophiles et d'un nom-
bre croissant d'eosinophiles. Le metoestrus se
caracterisait par la presence de cellules inter-
mediaires et parabasales, de neutrophiles et de
debris cellulaires. L'anoestrus se caracterisait par
la presence de groupes de cellules intermediaires et
de quelques cellules parabasales.
Au cours de cette etude, deux chattes demeure-
rent en anoestrus. On observa huit periodes
oestrales d'une duree de deux A cinq jours chacune,
chez les cinq chattes qui present&rent un cycle
normal. La duree du cycle oestral varia de 15 A 17
jours, chez trois chattes normales. Deux chattes ne
revinrent pas en oestrus au cours de l'experience,
mais l'examen bacteriologique d'ecouvillons vagi-
naux de ces deux chattes donna des resultats
positifs. Ceci n'exclut toutefois pas la possibilite
que leur oestrus du debut de l'experience se soit
accompagne d'une ovulation. Les frottis vaginaux
obtenus de ces deux chattes contenaient des
hematies, vraisemblablement reliees A l'infection
dont elles souffraient et qui s'accompagnait d'un
ecoulement vulvaire.
I N T R O D U C T IO N
The cyclical changes in exfoliative vaginal cytology
in the bitch have been extensively studied (5, 6) and
are widely used clinically. Only a few reports exist
on vaginal cytology in the feline species. The
morphological changes of feline vaginal epi-
thelium during the various stages of the natural
estrus cycle (2, 4, 7) and in stilbestrol-induced
estrus (3) have been described.
The purpose of this study was to examine
exfoliated vaginal cells in the cat through the estrus
cycle to determine if cyclic changes could be found
using the Maturation Index (MI) (12). Matura-
tion Index is the percentage of parabasal to
intermediate to superficial vaginal epithelial cells.
This index has not previously been described for
feline vaginal cytology. The cyclical changes were
supported by measurement of Eosinophilic Index
(El) and Karyopyknotic Index (KPI).
This cytological technique used in the cat will aid
in the management of reproductive failure, genital
University of Guelph, Guelph, Ontario NIG 2WI.
95
tract infections and in the optimal timing of
matings.
MATERIALS AND METHODS
Seven entire female cats aged between eight
months and five years were used in this study. The
group included a proven queen and three known
nulliparous cats. Vaginal smears were obtained
from each animal on alternate days for 32 days
during the months of May and early June (Spring).
The cats were housed in a controlled environment
at 200C, with 10.5 hours of artificial light per day.
The queens were individually caged in a 12 x 14 ft.
room also housing male cats, but no matings were
allowed during this study. The determination of
clinical estrus in each cat was based on behav-
ioural changes observed during and immediately
after each occasion of sampling. Behavioural
estrus (7) was measured by a subjective grading of 0
to 5. A score of 5 was used to indicate the most
marked clinical estrus with fast rolling, tail
deviation, elevation of hind quarters and treading.
A score of 0 indicated no change in behaviour in
the cat, and scores from I to 4 indicated increasing
intensities of clinical estrus. Behavioural estrus
scores were compared to the vaginal cytology
obtained.
Smooth glass rods (3 mm diameter and 10 cm
long) with smooth rounded ends were used for cell
collection. The rods were moistened with sterile
noted. The Eosinophilic Index (El) and Karyo-
pyknotic Index (KPI) for each specimen were
evaluated using Papanicolaou stained smears. The
El represents the ratio of all mature squamous cells
with eosinophilic cytoplasm to the mature squa-
mous cells with cyanophilic cytoplasm (12). The
KPI represents the ratio of all mature squamous
cells containing a pyknotic nucleus to mature
squamous cells containing a vesicular nucleus (12).
One month after the last serial sample was
obtained, vaginal swabs were taken from each cat
for bacterial culture and cytological examination.
In addition, quantitative cytology was done on
50 cells from each of the three maturation stages of
epithelial cells (parabasal, intermediate and super-
ficial) from Wright's stained smears. The cells and
nuclear dimensions were measured using a Zeiss
ocular micrometer (I mm in 100 divisions) which at
X25 objective, 100 ocular divisions covered
400 ,um and at X40 objective, 100 ocular divisions
covered 250 ,um. The mean cell dimensions and
standard deviations (s.d) were calculated. The
nucleus to cytoplasmic (N/C) ratios were deter-
mined using the area of cells calculated by length
times breadth as if the cells were rectangular.
RES U LTS
The epithelial cell types were readily recognized,
and their proportions in vaginal smears were
.-AL

normal saline and introduced into the vagina

approximately 1.5 cm, and quickly and gently
rotated against the floor and lateral walls of the
vagina. Two cellular spreads were immediately
prepared from each cat by rotating the rod along a
glass slide. One smear was air-dried and the other
was rapidly fixed with an aerosol cytological
fixative'. The air-dried slides were stained with
Wright's Romanowsky stain2. Alcohol spray-fixed
spreads were stained with a modified Papani-
colaou stain3.
The stained slides were examined at low mag-
nification to assess the cell yield, which was graded
as poor, moderate or good; and to assess the degree
of clumping of exfoliated cell populations which
was recorded as being grouped or single. Using
magnification of X160 and X400, 100 epithelial
cells on the Wright's stained smears were counted
and classified into groups of parabasal, inter-
mediate and superficial cells (I 1). Thus the,
Maturation Index (MI), the proportions of para-
basal to intermediate to superficial cells was
recorded for each specimen (12). The superficial
cell group was further subclassified into nucleated
and anucleated epithelial cells. The number of
neutrophils per 100 epithelial cells was counted
simultaneously. The presence of debris, bacteria FIGURE 1. Superficial feline vaginal epithelial cells at
and cytoplasmic vacuolation of epithelial cells was estrus. Wright's stain. Top X330, bottom X820.

'Pro-Fix, Lerner Laboratories, Stamford, Connecticut.

2Hema-Tek Slide Stainer, Ames Company, Elkhart, Indiana.
3Shandon-Elliot, Model SCE-0825, Shandon Southern Instruments Ltd., Surrey, England.

96
 TABLE I
DIMENSIONS OF FELINE VAGINAL EPITHELIAL CELLS (MICRONS)

Cytoplasm Nuclei
Cell Type length width length width N/C Ratio
Superficial 68.5 (l0.5)a 41.8 (11.9) 7.9 (1.2) 5.6 (1.5) 0.02
Intermediate 44.7 ( 8.9) 37.5 ( 7.7) 10.0 (2.1)b 0.06
Parabasal 18.6 ( 2.8)b 8.3 (1.7)b 0.20
aBrackets enclose one s.d. of mean
bNuclei round, single dimension given

applied in the characterization of phases of the

estrus cycle. Superficial epithelial cells were the
largest cells seen (Figure 1, Table I). They are oval
and flattened, with small and oval to pyknotic
nuclei and low N/C ratio. The morphology of the
intermediate epithelial cells is illustrated in Figure
2. These cells varied in size from small and
medium to large, but were not as large as
superficial cells. Intermediate cells were round to
oval in shape with smooth cytoplasmic and nuclear
membranes, and round and slightly flattened
vesicular nuclei. The parabasal epithelial cells
(Figure 3) were the smallest and thickest cells, with
a high N/C ratio and smooth, rounded cyto-
plasmic and nuclear membranes, and apparently
spherical nuclei. They were deeply cyanophilic and
were always grouped in layers of one to five cells
deep. The dimensions of the epithelial cell types are
given in Table I.

FIGURE 3. Parabasal feline vaginal epithelial cells.

...4. ....
FIGURE 2. Intermediate feline vaginal epithelial cells.
Wright's stain. Top X490, bottom X1070.
Wright's stain. Top (centre) X330, bottom X820.
Five cats showed estrus during the study, and
were given behavioural scores ranging from 2 to 5,
with a mean score of 4 on a total of 20 days of estrus
behaviour. The scores dropped at postestrus to a
mean of 1.9, ranging from 0 to 4. The remaining
two cats were in anestrus throughout the study and
had behavioural scores ranging from 0 to 3, with a
mean of 0.7. Scores tended to parallel cytological
findings of estrus.
The vaginal smears obtained from the five cats
exhibiting behavioural estrus were highly cellular
and free from debris. At estrus, epithelial cells were
shed singly and in high numbers. There were high
proportions of nucleated and anucleated super-
ficial epithelial cells, with a predominance of
nucleated superficial cells. Folding of cytoplasmic
borders was observed mainly in anucleate super-
ficial cells obtained in late estrus. Parabasal cells
and neutrophils were absent in estrus and inter-
mediate cells were rare. The superficial cell count

97
 TABLE II
THE MEANS AND RANGES OF PERCENTAGES OF PARABASAL, INTERMEDIATE SUPERFICIAL FELINE VAGINAL EPITHELIAL
CELLS. AND NEUTROPHILS. IN STAGES OF THE ESTRUS CYCLE

Neutrophils
Nucleated Anucleated per 100
No. of Parabasal Intermediate Superficial Superficial Epithelial
Cycle Stage Smears Cells Cells Cells Cells Cells

Estrus
Mean 20 0.3 11.6 63.6 24.5 4.7 d
Range 0-3 0-25 5-90 3-95 0-10
Early Metestrus a
Mean 8 8.9 75.7 13.2 1.9 32
Range 0-29 50-100 0-41 0-3 1-78
Late Metestrus
Mean 8 48 50 2 0 32
Range 10-85 10-85 0-5 0 0-97
A nestrus
Mean 34 9.7 87.4 2.7 0.2 3
Range 0-50 50-100 0-22 0-2 0-50
Proestrusc
Mean 4 17.8 60.3 19.6 2.2 8e
Range 0-34 57-67 6-40 0-6 0-33
aTwo days postestrus
"Four to nine days postestrus at time of maximum parabasal cells
'Two days before estrus
dNeutrophils only in estrus in three cats which also had bacteria on vaginal smears
'Neutrophils only in proestrus in one cat which also had bacteria on vaginal smears

averaged 88% in the eight estrus periods observed
and the mean percentage of parabasal to inter-
mediate to superficial epithelial cells (MI) in 20
smears of cats in estrus was 0:12:88 (Table II).
Anucleate superficial cells were only seen in estrus
and their numbers dropped off sharply to zero at
postestrus in the three cats which returned to estrus
during the study.
The behavioural estrus scores were variable
from period to period, and cytological changes
were found to be more reliable. Of these cyto-
logical changes, the MI was the most sharply
defined, followed by the KPI and the EI. At the
peak of behavioural estrus, the KPI and El were 90
to 100% in most of the cats. The El varied from 40
to 100% throughout estrus. Cytoplasmic eosino-
philia tended to precede by several days, the
changes in the KPI in early estrus, and the
eosinophilia persisted for two to four days longer
than nuclear pyknosis following estrus. The dura-
tion of the eight estrus periods observed was two to
five days.
During early netestrus (postestrus) the smears
contained mainly intermediate epithelial cells and
a low number of superficial cells. Cells in early
metestrus were moderately basophilic on Wright's
stain, and had irregular contours with mildly
pyknotic nuclei and cytoplasmic eosinophilia. The
mean MI in eight smears of five cats in early
metestrus at two days postestrus was 9:76:15
(Table II). Within four to nine days (average 6.4
days) after estrus, the presence of parabasal cells
increased to a mean peak of 48% (max. 85%), and
the El dropped to 10%. The mean MI in eight
smears in late metestrus was 48:50:2 (Table II).
Smears at this stage often contained debris, neu-
trophils and occasionally foam cells (Figure 4).
Neutrophils appeared after the last day of estrus in
seven of the eight estrus periods studied. The
highest proportions of neutrophils occurred at two
to eight days after the last day of estrus.
Vaginal smears obtained from the two cats in
anestrus tended to be poorly cellular, compared to
estrus smears, and had predominately medium-
sized intermediate cells usually grouped together,
and with a few parabasal cells. The mean MI of 34
smears of the cats in anestrus was 10:87:3 (Table
II). Two cats were behaviourally and cytologically
classified as being in anestrus throughout the
sampling period.
The smears obtained in proestrus were charac-
terized by high proportions of medium to large
intermediate cells and a few nucleated superficial
cells, with very little debris. The mean MI of four
smears obtained at proestrus was 18:60:22 (Table
II). Neutrophils were rare in these smears.
Erythrocytes were rarely seen and occurred only
with vaginal discharges in two cats which subse-
quently had bacterial growth on culture. Moderate
growths of hemolytic and nonhemolytic Staphylo-
cocci were cultured from the two cats, (P4 and P7)
which also showed bacteria on vaginal smears
during the study. The other five cats had no
significant bacterial growth.

98

FIGURE 4. Neutrophils in metestrus with intermediate
and parabasal vaginal epithelial cells. Several foam cells
are present on the left. Wright's stain. Top X330, bottom
X820.
The cycle lengths were 15, 17 and 17 days in three
cats, while behavioural estrus did not reappear in
two other cats during the 32-day study. Both of
these cats showed more irregular cellular yields,
with a mild persistence of anucleate superficial
cells following estrus. In addition, both cats had
bacterial growth on culture. An example of the
cyclical cell yield from one cat (P1) which showed
two estrus periods is given in Figure 5. The cellular
yields from a cat (P5) in anestrus. and from a cat
(P7) with one estrus period and vaginal bacteria on
culture, are shown in Figures 6 and 7 respectively.
THIN BAR - NEUTROPHILS PER 100 EPITHELIAL CELLS.
D I S C U S S IO N
The results from this preliminary study indicated
that the sequential estimation of the MI in vaginal
smears can be used in determining the phases of the
estrus cycle in the cat. Deviations from the normal
cyclic patterns of exfoliated cells may help to
indicate underlying reproductive problems such as
hormonal insufficiency or inflammation. Serum
reproductive hormone levels were not assayed in
this study but should be determined in future
studies in correlation with vaginal cytology. It has
been previously shown that vaginal cytology in the
cat reflects behavioural estrus (3), and this appear-
ed to be so in the present study.
The use of saline-moistened glass rods for
sample collection was atraumatic, preserved cell
integrity, and yielded more cells than moistened
cotton swabs used in a pilot study. The MI was
determined on Wright's stained smears, and this
stain appeared to provide adequate cytological
information for practical purposes. The Papan-
icolaou stained smears gave equivalent informa-
tion on the MI and allowed comparison with
findings in women. In contrast, the El determined
with Papanicolaou stain gave equivocal results
since some parabasal and small intermediate
epithelial cells had a degree of eosinophilia. The El
obtained, however, roughly paralleled behavioural
and cellular estrus. Variations in the Papanicolaou
stain reaction were related to cell thickness, stain
penetration, and to premature drying of the
preparations before alcohol fixation. Strasser et al
(10) also found variations with the Papanicolaou
stain in evaluating El in feline vaginal smears.
The Schorr Trichrome stain was tried in a pilot
study but was found to be unreliable for EI in our
hands. Eosinophilic Index is related to intra-
cellular factors and not to cell maturity ( 1). The
use of El in cytohormonal evaluation in women
has largely been abandoned in favour of the MI
which can be done with any stain, but usually with
Papanicolaou (12). The principle of the MI
determination is related to nuclear and cyto-
THICK BAR - VAGINAL EPITHELIAL CELLS. (ON LEFT AXIS)

P1
ESTRUS
BEHAVIOUR 4 55 0 0 1 3 3 3 2 3 5 2
4 0 0o
ANUCLEATED 100
SUPERFICIAL
NUCLEATED Notutrophils
SUPERFICIAL per 100
Epiethelial
INTERMEDIATE Ce' uIs

PARABASAL
I
I
I II, Il
lo
0 2 4 6 8 10 12 14 16 1. 20 22 24 26 28 30 32
TIME IN DAYS

FIGURE 5. The cyclical changes of exfoliated vaginal epithelial cell types in one cat (P1), which showed two periods of
clinical estrus.

99
 THIN BAR -
NEUTROPHILS PER 100 EPITHELIAL CELLS. THICK BAR - VAGINAL EPITHELIAL CELLS. (ON LEFT AXIS)

P5
ESTRUS
BEHAVIOUR 1 1 0 1 0 0 0 00 0 0 0 0 0 0 0
ANUCLEATED 100
SUPERFICIAL
NUCLEATED Neutrophils
SUPERFICIAL per 100
Epithelial
INTERMEDIATE Cells

PARABASAL i1111 w *

14
I

16 18 20 22 24 26 2 . 0
0 2 4 6 8 10 12 14 1,6 I'S 20 22 24 26 28 30 32
TIME IN DAYS
FIGURE 6. The cyclical changes of exfoliated vaginal epithelial cell types in one cat (P5) in anestrus throughout the study.

THIN BAR - NEUTROPHILS PER 100 EPITHELIAL CELLS. THICK BAR - VAGINAL EPITHELIAL CELLS. (ON LEFT AXIS)

P7
ESTRUS
BEHAVIOUR 2 0 0 0 0 0 1 1 1 2 3 2 1 1 1 2
MJCLEAED 100
SUPERIFICAL
IUCLEATED --A I Neutrophils
SUPERFICIAL per 100
Epithelial
I NTEMEDIATE Cells

PARABASAL I~~~~~~~~~I 0

0 2 4 6 8 10
ITIME IN DAYS
FIGURE 7. The cyclical changes of exfoliated vaginal epithelial cell types in one cat (P7) which had one estrus period and
hemolytic and nonhemolytic staphylococci on vaginal culture.
plasmic maturation of the epithelial cells as a
function of their size and thickness. These mor-
phological characteristics can be demonstrated by
any stain which gives good nuclear and cyto-
plasmic differentiation. Since blood stains are in
more general use than cytological stains, the
determination of MI by this method is most
practical. The MI is based on the maturation of the
cells and thus the tissue. On the other hand, the
KPI is more restricted to nuclear maturation and is
less informative than the MI.
The degree of folding of cytoplasmic margins
observed in these cats in estrus, is referred to as the
Folded Cell Index in women by Wied (12), and is
indicative of squamous cell maturation.
The cycle lengths of 15 and 17 days in these cats,
are consistent with the findings of previous
workers (4, 7). It is interesting that the two cats
which failed to show a recurrence of estrus within
30 days, also had vaginal bacteria on culture.
Ovulation in the cat depends on the neuro-
hormonal stimulation threshold and has been
induced by repeated vaginal stimulation using a
glass rod (1). It is possible that the collection
technique used in the present study may have
12 14 16 18 20 22 21 26 28 30 32
stimulated ovulation. Scott and Lloyd-Jacob (7)
found, however, that less than two consecutive
matings were insufficient to result in ovulation and
pregnancy. Estrus cycles in the unbred cat are
therefore anovular and corpora lutea are not
formed (9). As true metestrus with progesterone
release may not always occur, it is felt that the
period following anovular estrus in the unmated
cat may be more appropriately termed postestrus.
Serum progesterone levels could not be mea-
sured in this study to confirm hormonal status, so
it was not possible to differentiate ovular from
anovular estrus. The effect of estrogen on vaginal
epithelium can be readily assessed and reflects
ovarian activity (8), while the influence of pro-
gesterone is less lucid. The daylight period of 10.5
hours in the present experiment was out of our
control and may also have affected hormonal
release.
It is interesting that only the cats which cycled
showed high proportions of parabasal cells, usu-
ally following behavioural estrus and after the
exfoliation of mature superficial epithelial cells.
On the other hand, cats in anestrus consistently
exfoliated intermediate epithelial cells with only

100
rare parabasal cells. The irregular shedding of low
numbers of superficial epithelial cells in cat P7 for
12 days following estrus may be associated with the
finding of bacteria on vaginal smears at that time,
or could suggest irregular hormone balance. The
presence of bacterial vaginitis in women will cause
apparent epithelial cell eosinophilia and matura-
tion (I 1).
Neutrophil numbers and neutrophil inclusions
in epithelial cells were not found to the same extent
as in dogs in the early luteal phase (6), but the
presence of neutrophils may be related to the
hormone status and may be a difference between
ovular and anovular estrus. Foam cells were
occasionally seen in postestrus in this study. Feline
vaginal cytology differs from the bitch in lacking
erythrocytes, and it is interesting that the cats
which did show erythrocytes in their vaginal
smears also had bacterial growth on culture.
The MI is a useful index in reproductive
cytology, and is independent of stain variation and
is therefore a more reliable criteria than El or KPI
for cytohormonal evaluation. We anticipate that
vaginal cytology and cyclical patterns of cells
exfoliated will be increasingly used in clinical
situations of reproductive abnormalities in the cat
and will be most effective with serial sampling.
A C K N OW L E D G M E N T S
Thanks are due to Dr. B.J. McSherry, Depart-
ment of Pathology and to staff of the Clinical
Pathology section for excellent technical assis-
tance. The keen interest and advice from Dr.
W.T.K Bosu and Dr. R.C. Povey, Department of
Clinical Studies is gratefully acknowledged. We
are indebted to Dr. R.W. Gatehouse, Department
of Psychology for the use of his cats.
GAINES VETERINARY AWARD
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Limited, through Gaines Professional Service
Centre, has made available the "Gaines Veteri-
nary Award".
The award will be made to that veterinarian
whose work in small animal practice, clinical
research or basic sciences is judged to have
contributed significantly to the advancement of
small animal medicine, surgery or management
of small animal practice including advance-
ment of the public's knowledge of the respon-
sibilities of pet ownership.
Primary consideration shall be given to
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REFERENCES
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cat (Felis domestica). Anat. Rec. 58: 217-223. 1934.
2. LICHE. H. and K. WODZICKI. Vaginal smears and the
oestrous cycle of the cat and lioness. Nature, Lond.
144: 245-246. 1939.
3. MICHAEL, R.P. and P.P. scorr. The activation of
sexual behaviour in cats by the subcutaneous ad-
ministration of oestrogen. J. Physiol. 171: 254-
274. 1964.
4. MOWRER. R.T.. P.A. CONTI and C.F. ROSSOW. Vaginal
cytology: An approach to improvement of cat breed-
ing. Vet. Med. small Anim. Clin. 70: 691-696.
1975.
5. ROSZEL. J.F. Genital cytology of the bitch. Scope
XIX: 3-15. 1975.
6. SCHUTTE. A.P. Canine vaginal cytology. J. small
Anim. Pract. 8: 301-317. 1967.
7. scoTT. P.P. and M.A. LLOYD-JACOB. Studies on
Fertility. Volume VII. pp. 123-129. R.G. Harrison,
Editor. Oxford: Blackwell Scientific Publications.
1955.
8 SCOTT. P.P. The Cat. (Chap. 3). UFAW Handbook
on the Care and Management of Laboratory Ani-
mals. 3rd Edition. New York: Animal Welfare
Institute. 1967.
9. scoTr. P.P. Cats (Chap. 10). Reproduction and
Breeding Techniques on Laboratory Animals. pp.
192-208. E.S.E. Hafez, Editor. Philadelphia: Lea &
Febiger. 1970.
10. STRASSER, VON H.. R. BRUNK and C. BAEDER. Unter-
suchungen zum Sexualyklus der Katze. Berl. Munch.
tieirrztl. Wschr. 84: 253-254. 1971.
11. WIED, G.L. (Editor). The Symposia of the Inter-
national Academy of Gynecological Cytology. Acta
Cytol. 2. No. 1. 1958.
12. WIED, G.L. (Moderator). Symposium on Hormonal
Cytology. Acta Cytol. 12. No. 2. pp. 87-127. 1968.
Nominations for the 1979 Award may be made
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shall include a description of the work done by
the one nominated, a statement of how the work
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information.
The Award will consist of a gold medallion and
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All communications may be addressed to the
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