THE

AMERICAN

JOURNAL

OF

CLINICAL

NUTRITION

vol.

22, No. 4,

April,

1969,

pp.

391-395

Prinfr4in

U.S.A.

Intestinal Laying
i

Calcium Fowl and

1
#{149}

Absorption Its Importance

123

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

of the interest higher of

laying due to than the body

This its to in the

discussion features
OF

will of calcium

be

devoted absorption.

mainly

Downloaded from www.ajcn.org by guest on October 8, 2011

most other The total the laying

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).

methods The latter polydipsia lead to

for

exteriorizing methods result in

the in coloswater

increased changes take calcium 25-30 calcium absorption in

with place: (1),

ovar-polyuria and retomy does not metabolism from technique in

(7), whereas disturbances the studies. in laying (8).

were

and is therefore nutrition-balance were performed egg surgery

opening,

preferred several was Feces,

collected

about

mg/100 is

Almost all tein bound b) Increase

ml. Colostomies prohens. In (3). interrupted

tained from

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

the bone the appearance

(4). in

plastic

jars

and

the

urine

was

collected

formation

Since

chicken amounts per greater day,

weighs to a rate that than

less about that of

than 1,000 is a only absorbed The to about at

kg, mg least

the

activity of

of

the

organ is capable at a rate exceeding

1

transporting 100 mg/hr.

of Animal

uterus. Thisabsorption calcium calcium/kg 30

Science,

times

lactating about calurinary amount 10%

From

the

Department

woman. The

Since

the in

yolk most the similar

contains of in the eggshell. absolute only calcium. changes

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.

calcium, Rehovot, 25 mg cium is secreted while of man,

amounts absorbed purposes,

Department

of Agriculture,

Senior

of for

the total practical

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

Retentionb Loss in egg

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

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intake

total
-

excretion. loss in egg.

Retention

is (intake
reflect

The about

length 25-30 during

of

the laying cycle hr, with the shell the

need

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

The p1orus calcium,

requirement is considerably as the shell

of

the

chicken than mostly

for

phos-ject

variations.

lower contains

carbonate Over 90% in the yolk.

and only of the egg It is of

traces of phosphorus interest to

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

that calcium need must for be

cannot meet the and bone calcium

postulated that

most of the excreted the feces, whereas phosphorus appears

RELATIONSHIP BETWEEN

calcium is most of the in the urine.

CALCIUM

lized. He found in bone excreted from be regained

INTAKE

the

calcium

due to during

eggshell the

formation time when

would no egg-

AND

CALCIUM

ABSORPTION

Typical function ii. The

results of data intake were

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

calculation remained cycle. Stud(5) indicated the calexfor

the into

are presented obtained with substance,

a nonabsorbed

reference
TABLE

Table little aid laying of yttrium cium tent

shell

bone calcium during suggested that bone to

have

mobilized extra
must

a significant needed
come

calcium calcium elapse and excretion, to

II

formation

Calcium
in

absorption
hens during

as shell

a function formation
Ca absorption

of

body by intake Because

increase about 6 and not in a

in hr

absorption. between because balance

calof

cium intake the discontinuity

% of intake

excretion in fecal suitable absorption. nonabsorbed

study

Dietary

Ca,

Ca

intake

mg/day

Absolute, mg/day

methods are term changes searched for

to help stance

evaluate shortWe, therefore, reference

in

subcalcium

0.59 1.76 3.94

495 1,715 3,920

400 1,310 2,430

80.8
76.3 62.0

variations

absorption It was absorbed

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

Absorption the same

therefore

and that
be Percentage

Homeostasis
TABLE

393
III

was
can

as

of
hens,
formation

net
as
and

calcium
influenced
dietary

absorption
by
calcium

in

a suitable estimation of level of the lots of 3.56%

reference calcium intestine. diets The

substance absorption Then containlevel can

laying

shell

we
Dietary Ca,

studied two ing 1.90 and 3.56% cium is requirement, as

hens, fed calcium. the whereas

of calbe

Shell calcification
3.56 1.90

sligl1tly a were

above moderately uniformly

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

on the killed into each

in- Measured segrole played were

with

the

aid

by

the

intestine

in

calcium when the

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calcium at any from were

testine was Ca/#{176}Y ratios. In early shell

was

homeostasis in the laying hen. 91. as would be expected, in- However, was deficient in calcium, apparent diet mum absorption was inadequate

even those

maxito meet circumfed losses

each

lot,

shell calcification, calcification, and

being formed.

some during others when no

calcium need. Under during the stances this need was met late calcium, as bones shellof skeletal such a diet exhibited large

by

mobilization from hens calcium

The lated

overall from the

calcium absorption, Ca/#{176}Yratio in the

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

result due to having rate intake. in calearly

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

This finding by Bronsch and that theabsorpof calcium 45 was

has Since sulted cotion see alif

the in
(see intestinal

moderate some increase

I),

calcium in it by was the

method look for any were

depletion calcium of interest

could

reabsorpto
be

also Table

dose higher than

calcium

absorption

three being

when the eggshell further at the time whenacute

the

stimulated calcium
classical not balance

challenge For this

was rapid fed diets

of purpose,
sufficient, changes

an

deficiency.

process If one
into
is

take place. a constant (in

feed

flow
is accumulated

of

trients this

our

experience

nusince we did in absorption

Two groups

(13).
of hens con-

achieved

in the periods),

crop

during the it is evident shell intestinal

early evening that the formation absorption.

calcium during by increased This existence calcium

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

6 days. by a 2all birds calon were placed days 6

calcium-free the
for

(0.12%

experiment of an absorption

clearly hour-to-hour and the

demonstrates regulation very

the cium). Thereafter, their original diets of (repletion period). important

were
additional

Calcium

balances

394
TABLE
IV

Hurwitz

and mechanism different Using co-workers subjected transport

tinal intakes.

Bar of this response these two instances. is the same or

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

when appearance (17)

function

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

findings conducted periods. during the control and repletion

ab-

sorption Such

by an

the intestine extrapolation

the laying easy if

hen. one

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Results of this experiment dicate that both groups to calcium depletion by

in retention, from which an increase is likely in to during to rebuild largely

(Table IV) of hens responded a marked increase

to have absorption.

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

to support the overall these

at least

this conclusion and process of absorption. findings

to Harrison to constitute speculate

extend it is still

With for
mucosa

in of

mind the on

the birds relatively not quite value of

on the lower high absorption reached the 80%,

even though

calcium of probable the

intake, the nism calcium had maximum Harrison

the

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

function and during

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

J. Physiol. reproduction partition in pigeon

660, R., AND and calcium, plasma.

CALCIUM We

ABSORPTION

McDoNALD, the gen 1945.

TAYLOR,

M. of

0.
estrogen

The
administration

have in two
calcium

demonstrated calcium instances:

need

the homeostasis response

due to shell

role in to

of

the

phosphorus

intestine hens in
creased

laying an in3.

1.

Biol.
MOORE.

Chem.

T. and on

formation,

and response rivation. It

to the challenge is not clear,

of calcium however,

if

depthe

high phate
phosphorus

C., AND J. Ii. low levels of the prelaying

and on

dietary storage

The effect inorganic of calcium

of the

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.

Downloaded from www.ajcn.org by guest on October 8, 2011

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.