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AMERICAN
JOURNAL
OF
CLINICAL
NUTRITION
vol.
22, No. 4,
April,
1969,
pp.
391-395
Prinfr4in
U.S.A.
Intestinal Laying
i
in
the in
aicium
SHMUEL HURWITZ,
omeostasis
PH.D,4 AND A.
BAR, M.SC.5
HE
CALCIUM
METABOLISM
hen intensity,
is
of which
particular is much
discussion features
OF
will of calcium
be
devoted absorption.
mainly
of It is rather unfortunate for the investiegg gator is that, in birds, urine and feces are 2 g, most of which is in the shell. Since mixed a in the cloaca and excreted together; hen will lay nearly one egg per day, its caltherefore, surgical means are necessary in cium turnover rate will approach 0.1/day. order to separate these excretions. Available Such a high turnover rate requires exsurgical techniques include colostomy (6) hen is 20 g and that of one tremely efficient mechanisms for the assimi- and
ureters
animals calcium
studied. content
PARTITION
CALCIUM
EXCRETION
lation of calcium. During maturation, with ian activity, the following gard to calcium a)Increase 10 mg/lOO metabolism in plasma ml to the increased (2). in calcium by
various (7).
for
the in coloswater
about
mg/100 is
some by
the
cases, the
new
not ob-
c) Storage of calcium This is accompanied of medullary long bones. labile high d) involves and (5). The
in stainless steel trays placed under the wire bone in the cavities of the floor of the cage. Table I gives the balance The medullary bone is very sheet for both calcium and phosphorus its turnover rate is extremely from hens kept on a normal diet.
of the first eggshell
(4). in
plastic
jars
and
the
urine
was
collected
formation
Since
kg, mg least
the
activity of
of
the
times
From
the
Department
woman. The
Since
the in
Volcani Institute of Agricultural Research, Israel. Contribution from The Volcani Institute of excretion, Agricultural Research (N.U.I.A.) 1968, series no. E. to that
Supported States in Scientist. part by Research a grant Assistant. from under The PL United 480.
Department
of Agriculture,
Senior
of for
Therefore, in retention
391
392
TABLE
I
Hurwitz
and
91, from
Bar
hens during shell formation. The
Calcium
and
colostomized
phosphorus
hens
Calcium, g/day
balance
(8)
Phosphorus, g/day
of
experiments low. Calcium cium sorption However, sorption increase the highest
will absorption
be
described increased
in
detail with
ab-
becal-
Parameter
intake, whereas the percentage decreased with increased the decrease was small in in absolute intake
intake. abthe at
Intake
Fecal excretion
3.00
0.95
0.57 0. 18
in percentage comparison with and studied reached absorption a even absolute plateau. capacity level
Apparent
Urinary
absorption
excretion
2.05
0.22 1 .83 1.75
0.39
0.29 0.10 0.08 0.02
absorption
Bat ancec
a
0.08
-
absorption had not yet This indicates the high of the laying hen.
OF CALCIUM
Intake
Apparent
fecal
absorption
excretion.
-
urinary
excretion
REGULATION
ABSORPTION
intake
total
-
Retention
is (intake
reflect
The about
of
in
the being
is
hen desub-
minus
changes
total
in
excretion)
intestinal
would
absorption.
largely posited
the
last
19-20
for
hr.
Therefore,
physiological to large
calcium
of
the
for
phos-ject
variations.
lower contains
that for From calcium periments, shell phosphate. ing is note found sorption mineral, that
the
results
of (9)
calcium-balance
exdurabthis mobilost
Taylor formation
calculated intestinal
the
calcium
due to during
eggshell the
would no egg-
AND
CALCIUM
ABSORPTION
of
calcium
absorption in the
shell was being formed. This assumed that calcium absorption constant throughout the laying made by us with bones as ies a change cycle. was and not that an in This the
a nonabsorbed
reference
TABLE
mobilized extra
must
a significant needed
come
II
formation
Calcium
in
absorption
hens during
as shell
a function formation
Ca absorption
of
in hr
calof
Dietary
Ca,
Ca
intake
mg/day
Absolute, mg/day
subcalcium
80.8
76.3 62.0
variations
during the laying cycle. found (10) that yttrium 91 and that its rate of
was
not
#{176}
Measured
with
the
aid
of
Y.
passage
Calcium through of calcium considered for the at any the 45. intestine Yttrium
91
and that
be Percentage
Homeostasis
TABLE
393
III
was
can
as
of
hens,
formation
net
as
and
calcium
influenced
dietary
absorption
by
calcium
in
laying
shell
we
Dietary Ca,
of calbe
Shell calcification
3.56 1.90
sligl1tly a were
minimum 1.90% deficient labeled experimenand their various segment and yttrium level of the the killed
% Absorption
None
considered Both diets yttrium tal diets, testines ments. analyzed The net
diet. with
40.0
68.3
72.4
40.1
77.3
71.1
91y of
Early
Late
91. After 4 days the hens were were separated The contents for both absorption calculated some hens of
with
the
aid
by
the
intestine
in
homeostasis in the laying hen. 91. as would be expected, in- However, was deficient in calcium, apparent diet mum absorption was inadequate
even those
each
lot,
calcium need. Under during the stances this need was met late calcium, as bones shellof skeletal such a diet exhibited large
by
The lated
distal
testine, is given in Table III. As can be seen, calcium absorption shell formation increased markedly, almost Moreover, cium doubled there absorption at was both little levels difference periods of of
In time, such animals will also adapt calcu- (13). to the deficient diet by reducing their inlosses of calcium in the eggshell, with the of of thinner egg production
OF
AN
eggshells (14).
ACUTE
and
reduced
EFFECF
ON
DEPLETION
between
CALCIUM
ABSORPTION
and late shell calcification. been confirmed recently workers (12), who observed tion most was this of a single times formed did not assumes the intestine
because
the in
(see intestinal
reabsorpto
be
also Table
calcium
absorption
three being
stimulated calcium
classical not balance
of purpose,
sufficient, changes
an
deficiency.
process If one
into
is
flow
is accumulated
of
trients this
our
experience
(13).
of hens con-
achieved
in the periods),
crop
1.82 feeding taining This control need for day depletion can be met received a
and
3.92% calcium for period was followed period during which diet hens
an
calcium-free the
for
(0.12%
experiment of an absorption
were
additional
Calcium
balances
394
TABLE
IV
Hurwitz
Effect
of
2-day balance
calcium of laying
1.82
depletion hens
3.93
the on
(13)
in in
calcium
Dietary Ca,
vitro techniques, Kimberg and (1 5) showed that when rats were to a low calcium regimen, active of calcium developed in inteswhere no such transport was
Period
Control
Reple-
tion
Control
Repletion
segments Maddaiah
demonstrable
Ca Ca Ca Egg Ca Ca intake, excretion, retention, Ca, g/day g/day balance, retention, g/day g/day g/day 2.06 0.70 1 .36 1.56 -0.20 66.2 2.02 0.52 1 .51 1.38 0.13 74.6 4.13 2.34 1 .79 1.77 0.02 43.4
the
and
rats
Bronner
were
on
(16)
normal
showed
4.00
1 .96
2.04
1.43 0.61 51.3
that protein
inverse
the
of the
of
the
calcium-binding mucosa
calcium
in
intestinal
dietary
was
intake.
an these
It
is
most
to
tempting
the regulation
to of is
extrapolate
of calcium
ab-
sorption Such
by an
hen. one
evokes a carrier-mediated active transport. inHowever, results on the in vitro uptake of calcium by the intestinal mucosa of laying hens suggest simple diffusion as the mecharesulted nism of this transport (unpublished data). This,
In vivo studies ((18) and unpublished data)
together
enabled calcium lowing
with
the balance them
the
birds
decrease
in
maintain
shell
secretion,
a skeleton. thus
repletion, their
tend positive it to
alIn possible
extend it is still
With for
mucosa
in of
mind the on
regulation
and
calcium
(19) clearly a
mechaabsorption.
showed permeability
birds had barrier for calcium. Schachter (20) showed balance during that this permeability barrier in the rat acute short-term jejunum and ileum was oxygen dependent. depletion raised the calcium absorption to It thus seems possible that through some this higher level. This would suggest that metabolic change the permeability of the the response of the intestine to the chalmucosa may be modified and thus affect the lenge of calcium depletion is to some degree regulation of intestinal absorption. proportional to the magnitude of the chalbeen in a distinctly the control period, negative but an
lenge, the absorption of calcium being a
RIDDLE,
REFERENCES 0.,
physiology
calcium AND
of
the the
dietary experiment.
OF
calcium
fed
before 1.
W.
of
changes 76:
H.
in
REINHART.
Studies
in birds.
on
21.
the
reproduction
Blood
MECHANISM REGULATION OF
the
1926.
reproductive
RIDDLE.
cycle. effect
on and 159: 455, nitro-
Am. 2. of
CALCIUM We
ABSORPTION
M. of
0.
estrogen
The
administration
have in two
calcium
role in to
of
the
phosphorus
intestine hens in
creased
laying an in3.
1.
Biol.
MOORE.
Chem.
T. and on
formation,
of calcium however,
if
depthe
high phate
phosphorus
dietary storage
of phosand
composition
medullary
Calcium
and
35,
Absorption
Brit.
and
13.
Homeostasis
395
4.
5.
6.
7.
8.
9.
HuRwITz, S.,AND A. BAR. Calcium depletion and repletion in laying hens. 1. Effect on calcium in various bone segments, in egg shell and in HURWITZ, S. Calcium metabolism of pullets at blood plasma, and on calcium balance. Poultry the onset of egg production, as influenced by dietary calcium level. Poultry Sci. 43: 1462, 1964. Sci. 45: 345, 1966. 14. HURWITZ, S., AND P. GRIMINGER. Observations on HURWITZ, S. Bone composition and Ca5 retention in fowl as influenced by egg formation. Am. I. the calcium balance of laying hens. I. Agr. Sci. 54: 373, 1960. Physiol. 206: 198, 1964. 15. KIMBERG, D. V., D. ScHAcHTER, AND H. SCHENKER. ARIYOSHI, S., AND H. M0RIM0T0. Studies on the nitrogen metabolism in the fowl. 1. Separation Active transport of calcium by intestine: effect of dietary calcium. Am. I. Physiol. 200: 1256, of urine for nutritional balance studies. ull. B Natl. Inst. Agr. Sci.12: 37, 1956. 1961. 16. MADDAIAH, V. T., AND F. BRONNER. Calcium STURKIE, P. D., AND W. P. JoINER. Effects of cloacal cannulation on feed and water consumptransport and calcium binding protein (CaBP) tion in chickens. Poultry Sci. 38: 30, 1959. of rats. Symposium on Biophysical Aspects of HURWITZ, S., AND P. GRIMINGER. Partition of Permeability, Jerusalem, 1968, p. 18. calcium and phosphorus excretion in the laying 17. TAYLOR, A. N., AND R. H. WASSERMAN. Vitamin hen. Nature 189: 759, 1961. D,-induced calcium binding protein; partial TAYLOR, T. C. Calcium absorption and metabopurification, electrophoretic visualization, and lism in the laying hen. In: Nutrition of Pigs tissue distribution. Arch. Biochem. Biophys. 119: and Poultry, edited by J. T. Morgan and D.
in
cortical
bone
pullets.
J.
Nutr.
12:
1958.
536,
1967.
S., AND A. BAR. Activity, concentration, and lumen-blood electrochemical potential difcalcium-45 and yttrium-9l along the intestine, ference of calcium in the intestine of the laying and calcium absorption in the laying fowl. I. hen. J. Nutr. 95: 647, 1968. Nutr. 89: 311, 1965. 19. HARRISON, H. E., AND H. C. HARRISON. Vitamin 11. HURWITZ, S., AND A. BAR. Absorption of calcium D and permeability of intestinal mucosa to and phosphorus along the gastrointestinal tract calcium. Am. I. Physiol. 208: 370, 1965. of the laying fowl as influenced by dietary 20. SCHACHTER, D. Vitamin D and the active transcalcium and egg shell formation. Nutr. I. 86: port of calcium by the small intestine. The In: 433, 1965.
10. HuRwITz, S., AND
Lewis.
London:
Butterworths,
1961,
p.
148.
A.
BAR.
Rate
of
passage
of
18. HURWITZ,
12.
BRONSCH,
K., 14:
K. 105,
LORCHER
AND
B.
STADLER.
Zum
Veteri-
Transfer Biological
of
Calcium edited
and
Strontium
Across
Calciumstoffwechsel
ndrmed.
des
1967.
Geflugels.
Z.
Membranes,
man.
New
York:
Academic,
by R. H. Wasser1963, p. 197.