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Vitamin A in the Liver and Kidney of Some Felidae
VITAMIN A IN THE LIVER AND KIDNEY OF SOME FELIDAE BY RALPH HEYWOOD* North of England Zoological Society, Zoological Gardens, Chester
SUMMARY Liver and kidney vitamin A reserves offorty-one Felidae in a Zoological garden have been recorded. Liver vitamin A reserves were low in enteric disease. Storage of large quantities of vitamin A in the kidney is not characteristic of all Felidae and only in the kidneys ofjungle cat and caracallynx were appreciable quantities found. The presence of fat in the renal tubules was confirmed. There was no excretion of vitamin A in urines examined nor any evidence of large quantities of fat being excreted, save in the tiger. It is suggested that when a meat diet is fed, with the calcium/phosphorus ratio corrected to I: I, 200 i. u. of vitamin A per kg. provides for adequate liver storage. Vitamin A deficiency in lion cubs is discussed. INTRODUCTION Vitamin A deficiency in domestic cats was investigated by Geishoff, Andrus, Hegsted & Lentin (I957) and Scott and co-workers reported on aspects of feline nutrition including vitamin A (Greaves, Scott & Scott, I958; Scott, I96o; Scott & Greaves, I964; Scott, I965). Hypervitaminosis A has been studied in the cat and its association with deforming cervical spondylosis defined (Seawright, English & Gartner, I967). Large amounts of vitamin A were found to be stored in the kidney of the cat by Lowe, Morton & Vernon (I 95 7) who found a mean concentration of 98 i.u. Jg. in nine cats, which is higher than in livers of other species. The kidney vitamin A level was also investigated by Moore, Sharman & Scott (I963) and a higher mean reserve (ISO i.u.fg. kidney) was found in eighty-two cats examined. These authors demonstrated by fluorescence microscopy the presence of highly fluorescent fat globules due to vitamin A in certain parts of the kidney. These results were confirmed by Seawright et al. (I967). Little seems to be known about the vitamin A status of other Felidae. Liver oils from two lions, a tiger, a leopard, a racoon and a seal were reported to be devoid of vitamin A (Karrer, Euler & Schopp, I932) but these results may be suspect since the authors found only a trace of vitamin A in the liver oil of a polar bear, the liver of which is known to contain massive quantities of the vitamin (Rodahl, I949). The writer found reserves of 40 and 45 i.u.fg. in the
* Present address: 6 I nee Road, Thornton, Liverpool 23.
VITAMIN A IN FELIDAE
39 1
livers of twin stillborn polar bears. A reserve of 260 i. u.fg. ofliver was reported from a lion living for 20 years in a zoo (Gillam, 1938). The presence of fat globules in the renal tubules of cats is well documented (Moore et al., 1963). Fat globules were also observed in the proximal convoluted tubules of lion, tiger,cheetah and ocelot (Hewer, Harrison Matthews & Malkin, 1948). In the tiger, fat was noted in the urine during life and post mortem and it was estimated that up to 20 g. offat was excreted daily by both male and female. Stranack ( 1962) investigated feline kidneys by microdissection and found that the kidneys of tiger, lion, leopard, jaguar, cheetah, serval, fishing cat and domestic cat showed tubular fat. The distribution varied in different species and among individuals of the same species but not among individual nephrons within the same kidney; the droplet size varied. This paper presents the results of a vitamin A storage survey of Felidae dying at Chester Zoo. MATERIALS AND METHODS All cats dying in the gardens during the years 1964- 1966 were subjected to a routine post mortem, often within one hour of death. The liver and kidneys were weighed and samples taken for vitamin estimation. The vitamin was estimated by the antimony trichloride method, using an E.E.L. colorimeter. Extraction of the vitamin was by the method given by Ames, Risley & Harris (1954). Most samples were checked by extracting the vitamin with petrol ether at 6o °C for go minutes in a Soxhlet apparatus. Vitamin A in urine was estimated by the method given by Lawrie, Moore & Rajagopal (1941 ). Part of the kidney tissue was fixed in formal calcium (Bakers solution) and sections cut with a freezing microtome and stained for fat with Sudan Black. RESULTS The details of forty-one Felidae are shown in Table I. Stained sections of kidney showed fat globules at the edge of the convoluted tubules in all species examined. Some tubules did not take the stain. The glomeruli and medullary collecting tubules were free of fat. Globule size was small, and in the cheetah and margay cats the globules were very fine. Much larger globules were found in the caracal lynx and the 5-month-old jungle cat than in other species examined. Fat was present in the kidney of a newborn lion. These findings confirm those of Hewer et al. ( 1948) and Stranack ( 1961). The urines of two pumas, a lion, a clouded leopard, a margay cat and a jungle cat were negative for vitamin A. DISCUSSION It will be noted that most deaths were due to feline enteritis, which has been endemic in the gardens for some years. A vigorous vaccination programme is bringing the disease under control, though some deaths have been experienced in vaccinated stock. The disease takes one of three courses: ( 1) peracute, with death within hours of the first symptoms; (2) acute, with death 2 days after the
TABLE
Species
Cheetah Acinonyx jubatus Leopard Panthera pardus Leopard Panthera pardus (Black) Leopard Panthera pardus (Black) Leopard Panthera pardus Puma Felis concolor Puma Felis concolor Puma Felis concolor Puma Felis concolor Puma Felis concolor Clouded leopard Neofelis nebulosa Clouded leopard Neofelis nebulosa Margay cat Felis weidi Margay cat Felis weidi Caracal lynx Felis caracal Caracal lynx Felis caracal Caracal lynx Felis caracal Caracal lynx Felis caracal
Age (years)
Sex
Cause of death
I
"t.>:l
Liver weight (g .)
Liver vitamin A (i.u.Jg.)
Kidney weight (g.)
Kidney vitamin A (i.u.Jg.)
.. .:>
Ratio bodyweight to liver weight
Fat in proximal convoluted tubules
M
14
Cirrhosis of liver
648
1220
52'5
10
35: 1
+fine droplets
F
18
917'5
IO
63
Trace
-
M
5/I2
Hepatitis, chronic nephritis, ill for I month Chronic enteritis, vaccinated, vitamin A therapy given
-
480
-
Trace
-
++small droplets + +small droplets
Trace
-
Trace
-
+
24'5
5
7'7
Nil
2I :I
No fat seen
970
-
I6
-
Euthanasia
-
6Io
-
17
-
9/12
Peracute feline enteritis
493'5
IO
52'5
Trace
42:1
F
9/12
Peracute feline enteritis
485
8
47'5
Trace
37'5:1
M
IO/I2
Peracute feline enteritis
687
520
67
4
-
+very small droplets +small droplets +small droplets +small droplets Not examined
M
I I/12
Peracute feline enteritis
304
I280
42
4
-
F
IO/I2
Acute feline enteritis
248
IOO
38
17'5
27•8:1
F
I
Peracute feline enteritis
143'5
1260
14
8
22'3:1
11/12
Acute feline enteritis
139'6
240
14'4
4
22·6:1
F
I
Euthanasia, rickets
177
10
29
9
-
F
I0/12
Euthanasia, rickets
150
Trace
I9'5
Nil
33'2 :1
M
10/12
Acute feline enteritis
179
400
27'3
6J
-
++medium and small droplets ++ +medium and small droplets + fine droplets + fine droplets +++ large droplets ++large droplets Not examined
M
4/12
Chronic feline enteritis, vaccinated
91
Trace
13'5
7
32:1
Not examined
5/I2
M F
Full term foetus 20
Euthanasia
F
2
F
M
-
--~ >-l
( Jl
-
Chronic enteritis, vaccinated
F
td ~
small droplets
::t::
~
>-l ttl ~
><:
t_,
0
e z~
~~ ~
.:>
~
Age (years)
Liver weight (g, )
Liver vitamin A (i,u,/g,)
Kidney weight (g ,)
Kidney vitamin A (i,u,/g, )
Ratio bodyweight to liver weight
Fat in proximal convoluted tubules
Species
Sex
Caracal lynx Felis caracal Jungle cat Felis chaus Jungle cat Felis chaus Jungle cat Felis ehaus Jungle cat Felis ehaus Jungle cat Felis ehaus Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Pan/hera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion P anthera leo
M
4/12
Acute feline enteritis
100
10
13
15
F
5/ 12
Feline enteritis, vaccinated
110
20
15' 1
68
M
7 weeks
15
7 1'S
4' 2
7
M
7 weeks
20
72
4'1
8'5
27 '8 :1
Not examined
20,8
14 0
4'3
17
23: 1
Not examined
10'4
45
3'2
II
22:1
+small droplets + +small
F
8 weeks
F
1/12
F
9
M F M
Sti11born
M
Sti1lborn Stillborn
M
Stillborn
M
Stillborn
Cause of death
} Kill'" by mooha Accident Post parturition
2,200
d roplets Not examined
Nil
}p."n, .",.1 ~pu Ductus arteriosus
58
Nil Nil Nil
10'4 9,8
28'2:1 26:1
Not examined Not examined
5 6 'S
Nil
10'5
21,8:1
+small
14' 2
F
Liveborn
61
2'5 30 3'5 4 4 5 4
14
F F
14 days
~ Ea'ffi by mooha Protracted labour Deserted } Deserted
~
a:: ...... Z > ......
Z
t<:I
40 42'S
Nil
M F
Not examined
<: ......
":I
86,S
M
26,8:1
17
Liveborn
Liveborn Liveborn Stillborn Stillborn 1 day
+small droplets + + large
droplets Not examined
8
F F M F
35: 1
55'5 84'6 52
II
13
13'4 14 10,8 13,6 9'S
Nil
Trace 2 Nil Nil Trace
17'7 : 1
droplets Not examined
24'2:1
Not examined
32 :1 20:1 19' 2 :1
Not Not Not Not Not
t"" ...... t:i
~
examined examined examined examined examined
Not examiner! Not examined tAl
<.0 tAl
394
BRITISH VETERINARY JOURNAL, 123, 9
first symptoms and (3) chronic, usually met in vaccinated stock, when death occurs I week to a month after onset. It is to be expected that this disease, particularly in the chronic form, will lower the liver vitamin A reserve. This is illustrated in the clouded leopards and margay cats. The first animal infected showed the peracute form, and high vitamin A reserves were found; in the partner the disease took a more chronic course and much lower vitamin A reserves were found. The vitamin A status depends on dietary intake. Two caracallynx fed solely on horse meat, with no supplementary calcium or vitamin A, were destroyed because of rickets. Vitamin reserves were low. Two pumas dying in I965 of peracute feline enteritis had been fed a meat diet, supplemented with calcium to bring the calcium phosphorus ratio to I :I and with vitamin A at an estimated rate of IOO i.u. fkg. fday. These animals showed reserves of8 and IO i.u. jg. liver. The present practice is to give 200 i.u. jkg. fday and more adequate reserves are now found post mortem and the condition of the animals and their breeding ability have improved. This suggests that the cats have a higher requirement for vitamin A than other species. Vitamin A reserves in the kidneys were low save for one caracal and one jungle cat in which appreciable quantities were found. These findings are a marked contrast to those of Moore et al. ( I963) with kidneys of domestic cats. There was no correlation between the presence of fat in the renal tubules and vitamin A storage but the larger globules found in the caracal and jungle cat should however be further investigated in relation to presence of vitamin A. The presence of fat in the histological sections did not give positive evidence for fat excretion in large quantities in the urine of any of the species examined. A few fat globules were found in the urine of the lion and the clouded leopard. Clifford, Stowe & Good ( I 96 I ) recorded fat globules in the urine of one lioness examined. Observation on several species of cats living in the Zoo suggested that only in the tiger are large amounts of fat found in the urine. Clinical avitaminosis A has not been seen, but the condition was suspected in lion cubs. Cubs born during I964 were pale and still births occurred frequently. Anasarca of the fore quarters was a common feature. Of two cubs born in two different litters, one showed patent atrial septa and one a patent ductus arteriosus. No eye changes were found. Livers had a blotchy appearance and were friable. It was thought that the livers were rather large. The ratio of bodyweight to liver weight did not, however, indicate any real difference between normal cubs and those thought to be vitamin A deficient. Vitamin A estimations on the liver were negative, even when large quantities of liver were used. Liver vitamin A of newborn normal cubs was found to be 3·7 i.u.fg. (the mean of seven cubs). The level of 2 i.u. jg. is given for livers of newborn kittens (Van Eekelen & Wolff, cited by Moore, I957), though much higher levels were found in full term foetuses (Moore et al., I g63). A lioness died IO days after giving birth to vitamin A negative cubs, the liver reserves being 8 i.u.fg. This lioness showed marked thyriod hyperplasia. During the preceding 2 years she had been fed a diet solely of horse meat, and had given birth to three litters, none of which survived.
VITAMIN A IN FELIDAE
395
Stillborn lion cubs are frequently reported in zoological gardens (Urbain, I 95 I) and in the wild (Halloran, I 955). Specific vitamin A deficiency in lion cubs has been reported from Rotterdam Zoo (Van Bemmel, Zwart & Peters, I962). The occurrence of subcutaneous oedema and congenital anomalies of the heart, lung and mediastinum in thirty-two cubs born to two sister lionesses are recorded by Krediet & Zwart (I964). Giving dams 35,000 i.u. vitamin A daily brought about a marked improvement in breeding results and no further heart anomalies occurred. Rewell ( I 948) found three lion cubs that died at 2 days old had a patent ductus arteriosus and interatrial septum and thought the condition common in inbred lions. Anomalies recorded (Krediet, r 963; Krediet & Zwart, I964; Heywood, 1964) were not in inbred stock. The only congenital abnormality recorded in lions is cleft palate, which Heuschele ( r 959) thought not uncommon and due to dietary deficiency though genetic predisposition appeared necessary. ACKNOWLEDGMENTS
I wish to thank Mr. L. G. Cooper, Department of Anatomy, Liverpool University, for technical assistance in sectioning kidneys and for the photomicrographs. REFERENCES
AMEs, S. R., RisLEY, H. A. & liARrus, P. L. (1954). Ana(yt. Chern., 26, 1378. CLIFFORD, D. H ., SToWE, C. M. & GooD, A. C. (1961) . ] . Am. vet. med. Ass., 139, 111. GEISHOFF, S. N., ANDRUS, S. B., HEGSTED, D. M. & LENTIN, E. A. (1957) . Lab. Invest., 6, 227. GILLAM, A. E. (1938). Biochem. ]., 32, 1496. GREAVES, J. P ., ScoTT, P. P. & ScoTT, M. G. ( 1958). Proc. Nutr. Soc. Abstract 43· HALLORAN, P. O'CONNOR (1955). Am.]. vet. Res., 16, 61. HEUSCHELE, W. P. (1959). ]. Am. vet. med. Ass., 134, 365. HEWER, T. F., HARRISON MATTHEWS, L. & MALKIN, T. (1948). Proc. zoot. soc. Lond., n8, 924. KARRER, P., VAN EuLER, H. & ScHOPP, K. (1932). Helv. Chim. Acta, 15, 493· KREDIET, P. & ZwART, P. (1964). Tijdschr. Diergeneesk., 89 (suppl.) 32. LAwRIE, N. R ., MooRE, T. & RAJAGOPAL, K. R . (1941 ). Biochem. ]., 35, 825. LoWE, J. S., MoRTON, R. A. & VERNON, J. (1957). Biochem. ]., 67, 228-234. MooRE, T. (1957). Vitamin A. p. 242. Amsterdam and London : Elsevier Press . MooRE, T., SHARMAN, I. M. & ScoTT, P. P. (1963). R es. vet. Sci., 4, 397, 407. REWELL, R. E. (1948). Proc. zoot. Soc. Land., n8, 501. RoDAHL, K . (1949). Nature, Lond., 164, 530. ScoTT, P. P. (1960). Vet. R ec., 72, 8. ScoTT, P . P. & GREAVES, J.P. (1964). Proc. Nutr. Soc., 23, Abstract 34· ScoTT, P. P. (1965). Symposium on Canine and Feline Nutritional R equirements, p. 81 (ed . by 0 . Graham-] ones). Oxford: Pergamon Press. SEAWRIGHT, A. A., ENGLISH, P. B. & GARTNER, R. J. W. (1967). ]. comp. Path., 77, 29. STRANACK, F. (1962). Proc. zoot. Soc. Lond., 139, 375· URBAIN, A. (1951). Bull. Mus. Hist. nat., Paris, 23, 157· VAN BEMMEL, A. C. V., ZwART, P. & PETERS, J. C. (1962). Tijdschr. Diergeneesk. , 87, 826. (Accepted for publication 28th April 1967) La vita:rnine A dans le foie et les reins chez quelques £elides (Heywood) ResuiDe. Les reserves en vitamine A hepatique et renale chez 41 Felides d'unjardin zoologique ont ete enregistrees. Les reserves h epatiques etaient basses en cas de maladies enteriques. Le magasinage de grandes quantites de vitamine A n 'est pas characteristique de tousle Felides et
BRITISH VETERINARY JOUR NAL, I23, 9 des quantites appreciables dans les reins ont ete observees seulement chez le chat de Ia jungle et le caracal. La presence de gras dans le tubules renales a ete confirmee. Aucune evidence d'excretion de vitarnine A n'a ete observee dans les echantillons examines; egalement, aucune evidence n'a ete observee d'excretion de grandes quantites de gras, sauf chez le tigre. On suggere que lorsque !'alimentation consiste en viande et le rapport calcium/phosphorus est ajuste a I :I, 200 unites internationales suffisent pour un magasinage hepatique adequat. La carence de vitamine A chez les lionceaux est discutee.
VitaJDin A Gehalt der Leber und Nieren einiger katzenartiger Zootiere (Heywood) ZusaDlmenfassung. Der Vitamin A Gehalt der Leber und Nieren wurde in 4I Tieren der Felidae Familie eines zoologischen Gartens bestimmt. Darrnstorungen waren von einem Vitamin A Mangel begleitet. In der Regel gab es keine Speicherung des Vitamins in den Nieren von Felidae, und betriichtliche Mengen wurden nur in den Nieren von Dschungelkatzen und Caracal-Ltichsen gefunden. Gegenwart von Fett in den Nierenkaniilchen wurde b estiitigt. Im Harn wurde kein Vitamin A entdeckt, mit Ausnahme des Tigers enthielt der H arn auch kein Fett. Es wurde vorgeschlagen, dass in einer Fleischkost mit einem korrigierten Kalzium -Phosphorverhiiltnis von I :I, ein Vitamin A Gehalt von 200 I.E. pro kg. fur gentigende Leberspeicherung sorgt. Das Problem von Vitamin A Mangel in jungen Lowen wurde diskutiert.
La vitaDlina A en el higado y en los riii6nes en algunos felidos (Heywood) ResuDlen. Las reservas de vitamina A en el higado y riiion de 4I felidos en un jardin zool6gico han sido enregistradas. Las reservas hepaticas han sido bajas durante enfermedades entericas. El deposito de grandes cantidades de vitamina A en el riii6n no es caracteristico de todos los felidos y solo en los riii6nes del gato de Ia jungla y de Ia lince de Ia estepa han sido observadas cantidades apreciables. Se comproba Ia presencia de graso en los tubulos renales. En las muestras de orina examinadas no ha sido observada excrecion de vitamina Ani evidencia de importantes excreciones de graso, excepto en Ia tigra. Se sugiere que cuando el regimen alimenticio consiste en carne y la raz6n de calciojfosforo es ajustada a I: I, 200 unidades internacionales de vitamina A por kg. de peso corporal provee un deposito hepatico adecuado. Se discute Ia carencia de vitamina A en los cachorros de leon.
PLATE I
..
..!( .. .-.:•.
.. ~
.., F ig.
Fig.
Fig. 3·
Caracal lynx, X
1.
2.
Leopard,
X
290.
290.
Clouded leopard ,
X 290.
Heywood, Br. vet. ]. (1967), 123, 9
SUMMARY Liver and kidney vitamin A reserves offorty-one Felidae in a Zoological garden have been recorded. Liver vitamin A reserves were low in enteric disease. Storage of large quantities of vitamin A in the kidney is not characteristic of all Felidae and only in the kidneys ofjungle cat and caracallynx were appreciable quantities found. The presence of fat in the renal tubules was confirmed. There was no excretion of vitamin A in urines examined nor any evidence of large quantities of fat being excreted, save in the tiger. It is suggested that when a meat diet is fed, with the calcium/phosphorus ratio corrected to I: I, 200 i. u. of vitamin A per kg. provides for adequate liver storage. Vitamin A deficiency in lion cubs is discussed. INTRODUCTION Vitamin A deficiency in domestic cats was investigated by Geishoff, Andrus, Hegsted & Lentin (I957) and Scott and co-workers reported on aspects of feline nutrition including vitamin A (Greaves, Scott & Scott, I958; Scott, I96o; Scott & Greaves, I964; Scott, I965). Hypervitaminosis A has been studied in the cat and its association with deforming cervical spondylosis defined (Seawright, English & Gartner, I967). Large amounts of vitamin A were found to be stored in the kidney of the cat by Lowe, Morton & Vernon (I 95 7) who found a mean concentration of 98 i.u. Jg. in nine cats, which is higher than in livers of other species. The kidney vitamin A level was also investigated by Moore, Sharman & Scott (I963) and a higher mean reserve (ISO i.u.fg. kidney) was found in eighty-two cats examined. These authors demonstrated by fluorescence microscopy the presence of highly fluorescent fat globules due to vitamin A in certain parts of the kidney. These results were confirmed by Seawright et al. (I967). Little seems to be known about the vitamin A status of other Felidae. Liver oils from two lions, a tiger, a leopard, a racoon and a seal were reported to be devoid of vitamin A (Karrer, Euler & Schopp, I932) but these results may be suspect since the authors found only a trace of vitamin A in the liver oil of a polar bear, the liver of which is known to contain massive quantities of the vitamin (Rodahl, I949). The writer found reserves of 40 and 45 i.u.fg. in the
* Present address: 6 I nee Road, Thornton, Liverpool 23.
VITAMIN A IN FELIDAE
39 1
livers of twin stillborn polar bears. A reserve of 260 i. u.fg. ofliver was reported from a lion living for 20 years in a zoo (Gillam, 1938). The presence of fat globules in the renal tubules of cats is well documented (Moore et al., 1963). Fat globules were also observed in the proximal convoluted tubules of lion, tiger,cheetah and ocelot (Hewer, Harrison Matthews & Malkin, 1948). In the tiger, fat was noted in the urine during life and post mortem and it was estimated that up to 20 g. offat was excreted daily by both male and female. Stranack ( 1962) investigated feline kidneys by microdissection and found that the kidneys of tiger, lion, leopard, jaguar, cheetah, serval, fishing cat and domestic cat showed tubular fat. The distribution varied in different species and among individuals of the same species but not among individual nephrons within the same kidney; the droplet size varied. This paper presents the results of a vitamin A storage survey of Felidae dying at Chester Zoo. MATERIALS AND METHODS All cats dying in the gardens during the years 1964- 1966 were subjected to a routine post mortem, often within one hour of death. The liver and kidneys were weighed and samples taken for vitamin estimation. The vitamin was estimated by the antimony trichloride method, using an E.E.L. colorimeter. Extraction of the vitamin was by the method given by Ames, Risley & Harris (1954). Most samples were checked by extracting the vitamin with petrol ether at 6o °C for go minutes in a Soxhlet apparatus. Vitamin A in urine was estimated by the method given by Lawrie, Moore & Rajagopal (1941 ). Part of the kidney tissue was fixed in formal calcium (Bakers solution) and sections cut with a freezing microtome and stained for fat with Sudan Black. RESULTS The details of forty-one Felidae are shown in Table I. Stained sections of kidney showed fat globules at the edge of the convoluted tubules in all species examined. Some tubules did not take the stain. The glomeruli and medullary collecting tubules were free of fat. Globule size was small, and in the cheetah and margay cats the globules were very fine. Much larger globules were found in the caracal lynx and the 5-month-old jungle cat than in other species examined. Fat was present in the kidney of a newborn lion. These findings confirm those of Hewer et al. ( 1948) and Stranack ( 1961). The urines of two pumas, a lion, a clouded leopard, a margay cat and a jungle cat were negative for vitamin A. DISCUSSION It will be noted that most deaths were due to feline enteritis, which has been endemic in the gardens for some years. A vigorous vaccination programme is bringing the disease under control, though some deaths have been experienced in vaccinated stock. The disease takes one of three courses: ( 1) peracute, with death within hours of the first symptoms; (2) acute, with death 2 days after the
TABLE
Species
Cheetah Acinonyx jubatus Leopard Panthera pardus Leopard Panthera pardus (Black) Leopard Panthera pardus (Black) Leopard Panthera pardus Puma Felis concolor Puma Felis concolor Puma Felis concolor Puma Felis concolor Puma Felis concolor Clouded leopard Neofelis nebulosa Clouded leopard Neofelis nebulosa Margay cat Felis weidi Margay cat Felis weidi Caracal lynx Felis caracal Caracal lynx Felis caracal Caracal lynx Felis caracal Caracal lynx Felis caracal
Age (years)
Sex
Cause of death
I
"t.>:l
Liver weight (g .)
Liver vitamin A (i.u.Jg.)
Kidney weight (g.)
Kidney vitamin A (i.u.Jg.)
.. .:>
Ratio bodyweight to liver weight
Fat in proximal convoluted tubules
M
14
Cirrhosis of liver
648
1220
52'5
10
35: 1
+fine droplets
F
18
917'5
IO
63
Trace
-
M
5/I2
Hepatitis, chronic nephritis, ill for I month Chronic enteritis, vaccinated, vitamin A therapy given
-
480
-
Trace
-
++small droplets + +small droplets
Trace
-
Trace
-
+
24'5
5
7'7
Nil
2I :I
No fat seen
970
-
I6
-
Euthanasia
-
6Io
-
17
-
9/12
Peracute feline enteritis
493'5
IO
52'5
Trace
42:1
F
9/12
Peracute feline enteritis
485
8
47'5
Trace
37'5:1
M
IO/I2
Peracute feline enteritis
687
520
67
4
-
+very small droplets +small droplets +small droplets +small droplets Not examined
M
I I/12
Peracute feline enteritis
304
I280
42
4
-
F
IO/I2
Acute feline enteritis
248
IOO
38
17'5
27•8:1
F
I
Peracute feline enteritis
143'5
1260
14
8
22'3:1
11/12
Acute feline enteritis
139'6
240
14'4
4
22·6:1
F
I
Euthanasia, rickets
177
10
29
9
-
F
I0/12
Euthanasia, rickets
150
Trace
I9'5
Nil
33'2 :1
M
10/12
Acute feline enteritis
179
400
27'3
6J
-
++medium and small droplets ++ +medium and small droplets + fine droplets + fine droplets +++ large droplets ++large droplets Not examined
M
4/12
Chronic feline enteritis, vaccinated
91
Trace
13'5
7
32:1
Not examined
5/I2
M F
Full term foetus 20
Euthanasia
F
2
F
M
-
--~ >-l
( Jl
-
Chronic enteritis, vaccinated
F
td ~
small droplets
::t::
~
>-l ttl ~
><:
t_,
0
e z~
~~ ~
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~
Age (years)
Liver weight (g, )
Liver vitamin A (i,u,/g,)
Kidney weight (g ,)
Kidney vitamin A (i,u,/g, )
Ratio bodyweight to liver weight
Fat in proximal convoluted tubules
Species
Sex
Caracal lynx Felis caracal Jungle cat Felis chaus Jungle cat Felis chaus Jungle cat Felis ehaus Jungle cat Felis ehaus Jungle cat Felis ehaus Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Pan/hera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion Panthera leo Lion P anthera leo
M
4/12
Acute feline enteritis
100
10
13
15
F
5/ 12
Feline enteritis, vaccinated
110
20
15' 1
68
M
7 weeks
15
7 1'S
4' 2
7
M
7 weeks
20
72
4'1
8'5
27 '8 :1
Not examined
20,8
14 0
4'3
17
23: 1
Not examined
10'4
45
3'2
II
22:1
+small droplets + +small
F
8 weeks
F
1/12
F
9
M F M
Sti11born
M
Sti1lborn Stillborn
M
Stillborn
M
Stillborn
Cause of death
} Kill'" by mooha Accident Post parturition
2,200
d roplets Not examined
Nil
}p."n, .",.1 ~pu Ductus arteriosus
58
Nil Nil Nil
10'4 9,8
28'2:1 26:1
Not examined Not examined
5 6 'S
Nil
10'5
21,8:1
+small
14' 2
F
Liveborn
61
2'5 30 3'5 4 4 5 4
14
F F
14 days
~ Ea'ffi by mooha Protracted labour Deserted } Deserted
~
a:: ...... Z > ......
Z
t<:I
40 42'S
Nil
M F
Not examined
<: ......
":I
86,S
M
26,8:1
17
Liveborn
Liveborn Liveborn Stillborn Stillborn 1 day
+small droplets + + large
droplets Not examined
8
F F M F
35: 1
55'5 84'6 52
II
13
13'4 14 10,8 13,6 9'S
Nil
Trace 2 Nil Nil Trace
17'7 : 1
droplets Not examined
24'2:1
Not examined
32 :1 20:1 19' 2 :1
Not Not Not Not Not
t"" ...... t:i
~
examined examined examined examined examined
Not examiner! Not examined tAl
<.0 tAl
394
BRITISH VETERINARY JOURNAL, 123, 9
first symptoms and (3) chronic, usually met in vaccinated stock, when death occurs I week to a month after onset. It is to be expected that this disease, particularly in the chronic form, will lower the liver vitamin A reserve. This is illustrated in the clouded leopards and margay cats. The first animal infected showed the peracute form, and high vitamin A reserves were found; in the partner the disease took a more chronic course and much lower vitamin A reserves were found. The vitamin A status depends on dietary intake. Two caracallynx fed solely on horse meat, with no supplementary calcium or vitamin A, were destroyed because of rickets. Vitamin reserves were low. Two pumas dying in I965 of peracute feline enteritis had been fed a meat diet, supplemented with calcium to bring the calcium phosphorus ratio to I :I and with vitamin A at an estimated rate of IOO i.u. fkg. fday. These animals showed reserves of8 and IO i.u. jg. liver. The present practice is to give 200 i.u. jkg. fday and more adequate reserves are now found post mortem and the condition of the animals and their breeding ability have improved. This suggests that the cats have a higher requirement for vitamin A than other species. Vitamin A reserves in the kidneys were low save for one caracal and one jungle cat in which appreciable quantities were found. These findings are a marked contrast to those of Moore et al. ( I963) with kidneys of domestic cats. There was no correlation between the presence of fat in the renal tubules and vitamin A storage but the larger globules found in the caracal and jungle cat should however be further investigated in relation to presence of vitamin A. The presence of fat in the histological sections did not give positive evidence for fat excretion in large quantities in the urine of any of the species examined. A few fat globules were found in the urine of the lion and the clouded leopard. Clifford, Stowe & Good ( I 96 I ) recorded fat globules in the urine of one lioness examined. Observation on several species of cats living in the Zoo suggested that only in the tiger are large amounts of fat found in the urine. Clinical avitaminosis A has not been seen, but the condition was suspected in lion cubs. Cubs born during I964 were pale and still births occurred frequently. Anasarca of the fore quarters was a common feature. Of two cubs born in two different litters, one showed patent atrial septa and one a patent ductus arteriosus. No eye changes were found. Livers had a blotchy appearance and were friable. It was thought that the livers were rather large. The ratio of bodyweight to liver weight did not, however, indicate any real difference between normal cubs and those thought to be vitamin A deficient. Vitamin A estimations on the liver were negative, even when large quantities of liver were used. Liver vitamin A of newborn normal cubs was found to be 3·7 i.u.fg. (the mean of seven cubs). The level of 2 i.u. jg. is given for livers of newborn kittens (Van Eekelen & Wolff, cited by Moore, I957), though much higher levels were found in full term foetuses (Moore et al., I g63). A lioness died IO days after giving birth to vitamin A negative cubs, the liver reserves being 8 i.u.fg. This lioness showed marked thyriod hyperplasia. During the preceding 2 years she had been fed a diet solely of horse meat, and had given birth to three litters, none of which survived.
VITAMIN A IN FELIDAE
395
Stillborn lion cubs are frequently reported in zoological gardens (Urbain, I 95 I) and in the wild (Halloran, I 955). Specific vitamin A deficiency in lion cubs has been reported from Rotterdam Zoo (Van Bemmel, Zwart & Peters, I962). The occurrence of subcutaneous oedema and congenital anomalies of the heart, lung and mediastinum in thirty-two cubs born to two sister lionesses are recorded by Krediet & Zwart (I964). Giving dams 35,000 i.u. vitamin A daily brought about a marked improvement in breeding results and no further heart anomalies occurred. Rewell ( I 948) found three lion cubs that died at 2 days old had a patent ductus arteriosus and interatrial septum and thought the condition common in inbred lions. Anomalies recorded (Krediet, r 963; Krediet & Zwart, I964; Heywood, 1964) were not in inbred stock. The only congenital abnormality recorded in lions is cleft palate, which Heuschele ( r 959) thought not uncommon and due to dietary deficiency though genetic predisposition appeared necessary. ACKNOWLEDGMENTS
I wish to thank Mr. L. G. Cooper, Department of Anatomy, Liverpool University, for technical assistance in sectioning kidneys and for the photomicrographs. REFERENCES
AMEs, S. R., RisLEY, H. A. & liARrus, P. L. (1954). Ana(yt. Chern., 26, 1378. CLIFFORD, D. H ., SToWE, C. M. & GooD, A. C. (1961) . ] . Am. vet. med. Ass., 139, 111. GEISHOFF, S. N., ANDRUS, S. B., HEGSTED, D. M. & LENTIN, E. A. (1957) . Lab. Invest., 6, 227. GILLAM, A. E. (1938). Biochem. ]., 32, 1496. GREAVES, J. P ., ScoTT, P. P. & ScoTT, M. G. ( 1958). Proc. Nutr. Soc. Abstract 43· HALLORAN, P. O'CONNOR (1955). Am.]. vet. Res., 16, 61. HEUSCHELE, W. P. (1959). ]. Am. vet. med. Ass., 134, 365. HEWER, T. F., HARRISON MATTHEWS, L. & MALKIN, T. (1948). Proc. zoot. soc. Lond., n8, 924. KARRER, P., VAN EuLER, H. & ScHOPP, K. (1932). Helv. Chim. Acta, 15, 493· KREDIET, P. & ZwART, P. (1964). Tijdschr. Diergeneesk., 89 (suppl.) 32. LAwRIE, N. R ., MooRE, T. & RAJAGOPAL, K. R . (1941 ). Biochem. ]., 35, 825. LoWE, J. S., MoRTON, R. A. & VERNON, J. (1957). Biochem. ]., 67, 228-234. MooRE, T. (1957). Vitamin A. p. 242. Amsterdam and London : Elsevier Press . MooRE, T., SHARMAN, I. M. & ScoTT, P. P. (1963). R es. vet. Sci., 4, 397, 407. REWELL, R. E. (1948). Proc. zoot. Soc. Land., n8, 501. RoDAHL, K . (1949). Nature, Lond., 164, 530. ScoTT, P. P. (1960). Vet. R ec., 72, 8. ScoTT, P . P. & GREAVES, J.P. (1964). Proc. Nutr. Soc., 23, Abstract 34· ScoTT, P. P. (1965). Symposium on Canine and Feline Nutritional R equirements, p. 81 (ed . by 0 . Graham-] ones). Oxford: Pergamon Press. SEAWRIGHT, A. A., ENGLISH, P. B. & GARTNER, R. J. W. (1967). ]. comp. Path., 77, 29. STRANACK, F. (1962). Proc. zoot. Soc. Lond., 139, 375· URBAIN, A. (1951). Bull. Mus. Hist. nat., Paris, 23, 157· VAN BEMMEL, A. C. V., ZwART, P. & PETERS, J. C. (1962). Tijdschr. Diergeneesk. , 87, 826. (Accepted for publication 28th April 1967) La vita:rnine A dans le foie et les reins chez quelques £elides (Heywood) ResuiDe. Les reserves en vitamine A hepatique et renale chez 41 Felides d'unjardin zoologique ont ete enregistrees. Les reserves h epatiques etaient basses en cas de maladies enteriques. Le magasinage de grandes quantites de vitamine A n 'est pas characteristique de tousle Felides et
BRITISH VETERINARY JOUR NAL, I23, 9 des quantites appreciables dans les reins ont ete observees seulement chez le chat de Ia jungle et le caracal. La presence de gras dans le tubules renales a ete confirmee. Aucune evidence d'excretion de vitarnine A n'a ete observee dans les echantillons examines; egalement, aucune evidence n'a ete observee d'excretion de grandes quantites de gras, sauf chez le tigre. On suggere que lorsque !'alimentation consiste en viande et le rapport calcium/phosphorus est ajuste a I :I, 200 unites internationales suffisent pour un magasinage hepatique adequat. La carence de vitamine A chez les lionceaux est discutee.
VitaJDin A Gehalt der Leber und Nieren einiger katzenartiger Zootiere (Heywood) ZusaDlmenfassung. Der Vitamin A Gehalt der Leber und Nieren wurde in 4I Tieren der Felidae Familie eines zoologischen Gartens bestimmt. Darrnstorungen waren von einem Vitamin A Mangel begleitet. In der Regel gab es keine Speicherung des Vitamins in den Nieren von Felidae, und betriichtliche Mengen wurden nur in den Nieren von Dschungelkatzen und Caracal-Ltichsen gefunden. Gegenwart von Fett in den Nierenkaniilchen wurde b estiitigt. Im Harn wurde kein Vitamin A entdeckt, mit Ausnahme des Tigers enthielt der H arn auch kein Fett. Es wurde vorgeschlagen, dass in einer Fleischkost mit einem korrigierten Kalzium -Phosphorverhiiltnis von I :I, ein Vitamin A Gehalt von 200 I.E. pro kg. fur gentigende Leberspeicherung sorgt. Das Problem von Vitamin A Mangel in jungen Lowen wurde diskutiert.
La vitaDlina A en el higado y en los riii6nes en algunos felidos (Heywood) ResuDlen. Las reservas de vitamina A en el higado y riiion de 4I felidos en un jardin zool6gico han sido enregistradas. Las reservas hepaticas han sido bajas durante enfermedades entericas. El deposito de grandes cantidades de vitamina A en el riii6n no es caracteristico de todos los felidos y solo en los riii6nes del gato de Ia jungla y de Ia lince de Ia estepa han sido observadas cantidades apreciables. Se comproba Ia presencia de graso en los tubulos renales. En las muestras de orina examinadas no ha sido observada excrecion de vitamina Ani evidencia de importantes excreciones de graso, excepto en Ia tigra. Se sugiere que cuando el regimen alimenticio consiste en carne y la raz6n de calciojfosforo es ajustada a I: I, 200 unidades internacionales de vitamina A por kg. de peso corporal provee un deposito hepatico adecuado. Se discute Ia carencia de vitamina A en los cachorros de leon.
PLATE I
..
..!( .. .-.:•.
.. ~
.., F ig.
Fig.
Fig. 3·
Caracal lynx, X
1.
2.
Leopard,
X
290.
290.
Clouded leopard ,
X 290.
Heywood, Br. vet. ]. (1967), 123, 9