The Physiology of Induced Molting
W. D. Berry1
Department of Poultry Science, 232 Upchurch Building, Auburn University, Auburn, Alabama 36849
ABSTRACT The initiation of seasonal feather molting
in wild avian species frequently coincides with incubation
of eggs and brooding of offspring. A period of natural
inappetence or anorexia usually accompanies this molt.
This is particularly true of the jungle fowl, the wild ances-
tor of the domestic chicken. Brooding of eggs by the jungle
fowl is accompanied by spontaneous anorexia, with little
food or water consumed throughout the period of egg
incubation. During this time, the reproductive tract re-
gresses, and feather molting is initiated. Selective breed-
ing for a high rate of egg production has blunted the
response of the commercial laying hen to exogenous envi-
ronmental cues and reduced or eliminated the endoge-
nous biological cues that coordinate initiation of seasonal
molting. However, commercial layers retain in their phys-
(Key words: induced molting, molt, molting physiology, immune function, laying hen)
INTRODUCTION
Molting in avian species may be generally defined as
the periodic shedding and replacement of feathers. For
most wild species of birds, molting involves reproductive
quiescence. The domestic hen likewise experiences a de-
crease in reproductive function during a naturally oc-
curring molt. However, this reproductive quiescence has
been found to be incomplete, and the hen often continues
to lay eggs at a low rate for a prolonged period. Therefore,
for the commercial egg producer, there would be a period
of unprofitably low egg production signifying the end of
the useful life of a flock. To avoid this, most hens are sold
and replaced just prior to the onset of natural molting.
Decreasing prices for eggs and spent flocks generated
interest in methods by which the natural molt could be
avoided and flocks kept for more than 1 yr. “Forcing”
the onset of molting to occur other than at the time of the
natural molting completely halted reproductive function
and precipitated a loss of feathers. Egg production re-
sumed and increased rapidly to a profitable rate following
this artificial molt. Thus, the term “forced molting” was
2003 Poultry Science Association, Inc.
Received for publication October 18, 2002.
Accepted for publication March 19, 2003.
1
To whom correspondence should be addressed: berrywd@acesag.
auburn.edu.
971
iological repertoire the ability to tolerate prolonged fast-
ing and to undergo a spontaneous regression of the repro-
ductive tract and feather molting. Induction of a coordi-
nated molt, by manipulation of environmental and
nutritional cues, or endocrine manipulation, can be used
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in domestic hens to regress and regenerate the reproduc-
tive tract. This improves subsequent egg production and
eggshell quality. This process also induces temporary re-
crudescence of lymphoid tissues and may alter immune
function in hens. The process of molting, and the subse-
quent recovery from the molt, may be viewed as a com-
plex physiological constellation, induced by environmen-
tal and nutritional cues, involving endocrine systems, re-
productive tissue structure and function, lymphoid
structure, and immune function.
2003 Poultry Science 82:971–980
coined and came into common usage. Other terms such
as pause, rest, forced rest, and recycling have been used
synonymously with the term forced molting. However,
artificially molting is the induction of a response that is
a natural component of the physiology of the hen. For
this reason, induced molting has been suggested as the
technically correct term.
Conventional induced molting usually involves remov-
ing feed, water, or both from the hens and reducing the
photoperiod to that of natural day length or less. Hens
are then fasted for a length of time sufficient to affect
complete involution of the reproductive tract. If water
restriction is involved, it is usually removed at the begin-
ning of fasting and may be continued for up to 3 d.
Molting is said to be induced during this time, as egg
production ceases and the reproductive tract rapidly in-
volutes. The period during fasting and following refeed-
ing when egg production is essentially at zero is referred
to as the rest period. During the rest period following
fasting, various molting, developer, or resting rations are
often fed as a part of some molting programs. With the
onset of renewed egg production, a layer ration is re-
Abbreviation Key: GnRH = gonadotropin-releasing hormone; HPA
= hypothalamic-pituitary-adrenal; HPG = hypothalamic-pituitary-go-
nadal; LH = luteinizing hormone; (1,25(OH)2D3) = 1,25 dihydroxychole-
calciferol; RBC = red blood cells; T3 = triiodothyronine; T4 = thyroxin.
972 BERRY
turned to the flock, and a supplemental lighting program
is initiated. Egg production then rapidly increases to a
level above that immediately preceding the induced molt.
By this time rejuvenation has occurred. The length of
molting, defined as the time from initiation of fasting until
50% egg production, varies with the particular molting
technique and conditions. Induced molts generally range
from 5 to 9 wk in length (Swanson and Bell, 1974a,b,c;
Brake and Carey, 1983).
Various induced molting techniques that do not use
fasting have been suggested. These alternate methods of
induced molting use dietary manipulations to create an
imbalance of a particular nutrient or nutrients (Douglas
et al., 1972; Creger and Scott, 1977; Berry and Brake, 1985,
1987). Investigations into the feasibility of alternate molt-
ing techniques have been concerned only with the effects
of these methods on egg production and shell quality.
Only a few studies have investigated the physiological
effects of these alternate induced molting techniques or
compared alternative versus conventional induced molt-
ing (Berry and Brake, 1987, 1991; Berry et al., 1987).
Induced molting is an important tool for the economic
management of laying flocks. Concerns about the effects
of fasting on the welfare of the hens and reports that
suggest that molting hens may be more susceptible to
Salmonella infection have led to calls for the abolition of
induced molting. Curtailment of induced molting would
be expected to reduce the profitability of the laying hen
enterprise and increase the cost of eggs and egg products
to the consumer. Understanding the physiological effects
of molting is necessary to rationally address hen welfare
and food safety concerns and to most efficiently and hu-
manely manage laying flocks. This paper will compare
the induction of molting in wild and domestic species
and will examine the physiological changes that accom-
pany the induction and recovery from an induced molt
with regard to the endocrine, reproductive, and immune
and hematopoietic systems.
INDUCED MOLTING TECHNIQUES
Induced molting techniques have varied widely in
methods for initiating molting and halting egg produc-
tion. Conventional induced molting strategies have used
fasting of hens for a particular length of time (Swanson
and Bell, 1974b; Rolon et al., 1993), or to a targeted BW
(Brake and Carey, 1983), with or without water restriction.
Improved postmolt performance of flocks exposed to a
short photoperiod during molt (Hansen, 1966a, 1969) led
to inclusion of photoperiod modification in many conven-
tional induced molting procedures (Swanson and Bell,
1974b).
Alternate molting techniques have used various nutri-
tional imbalances to induce molting. Methods studied
included feeding a ration containing less than 0.3% Ca
(Douglas et al., 1972; Martin et al., 1973; Gilbert and Blair,
1975; Wakeling, 1978), feeding Na-deficient rations
(<0.04% Na) (Whitehead and Shannon, 1974; Dilworth
and Day, 1976; Nesbeth et al., 1976a,b; Campos and Baiao,
1979; Ross and Herrick, 1981), feeding a ration containing
high levels (20,000 mg/kg) of Zn as ZnO (Creger and
Scott, 1977), and feeding a diet containing 2,500 to 5,000
mg/kg I as potassium iodide (Arrington et al., 1967).
Conventional fasting techniques for induced molting
cause complete cessation of egg production within 10
d from initiation of fasting (Swanson and Bell, 1971).
Alternative molting techniques, with the exception of
high Zn diets, have not proven to be as consistent in
halting egg production.
The efficacy of low Ca diets to halt production varies
widely. Martin et al. (1973) found that a dietary Ca level
of 0.3% caused a complete cessation of lay within 14
d in 69-wk-old Leghorn hens. Gilbert and Blair (1975)
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observed that 0.05% dietary Ca reduced production to
less than 5% by 35 d in 32-wk-old pullets. A Ca level of
0.09% with 0.32% P decreased production from 55 to 3%
in 14 d in 10-mo-old pullets and decreased production
from 70 to 2% in 42 d in 6-mo-old pullets (Douglas et al.,
1972). However, after 3 d on the low Ca diet, many hens
exhibited paralysis.
Low Na rations for molting laying hens have received
considerable attention (Wakeling, 1977). Diets deficient in
Na inhibit egg production when administered to pullets
prior to egg laying (Whitehead and Shannon, 1974). Inves-
tigations into the use of low Na rations for induced molt-
ing, however, have met with limited success. Whitehead
and Shannon (1974) found that a diet containing 0.038%
Na caused egg production to decline from 50 to 0% within
4 wk in a flock of 70-wk-old hens. Hens fed a diet without
added Na for 28 d exhibited a decrease in egg production
from 74% to less than 10% in 4 wk (Dilworth and Day,
1976). Nesbeth et al. (1976a,b) reported that a diet calcu-
lated to contain 0.017% Na fed to 11-mo-old pullets re-
sulted in complete cessation of egg production in 21 d in
one trial but only reduced production to 12% in another
similar trial. Naber et al. (1980) halted production within
15 d with a low Na diet. However, special care was taken
in that each feed ingredient was individually analyzed
for Na content before inclusion into the experimental diet
(Naber et al., 1980). Ross and Herrick (1981) used a diet
calculated to contain 0.044% Na but which upon analysis
was found to contain 0.13% Na. They reported that pro-
duction decreased from 60 to 15% in 5 wk.
Dietary Zn, usually in the form of ZnO, is reported
to be an effective agent for halting egg production and
inducing molting. Zn, when fed at a level of 20,000 ppm,
caused a complete cessation of egg production within 5
d (Scott and Creger, 1976; Creger and Scott, 1977; Berry
and Brake, 1985, 1987). Roberson and Francis (1979) re-
ported that 20,000 ppm dietary Zn is as effective as fasting
for inducing molting. Shippee et al. (1979) observed that
10,000 mg/kg Zn as oxide or acetate causes production
to decline from 60 to 0% in 6 d. Combining a low calcium
diet with a relatively low level of Zn (2,800 ppm) induces
reproductive involution and molting similar to that of
20,000 ppm dietary Zn (Breeding et al., 1992).
High dietary levels of I (iodine) were found to induce
a pause in egg production with or without loss of feathers.
 SYMPOSIUM: INDUCED MOLTING IN LAYERS
Arrington et al. (1967) observed that feeding 2,500 to 5,000
mg/kg I as potassium iodide to hens resulted in complete
cessation of egg production within 1 wk. Hebert and
Cerniglia (1979) reported that high levels of I stopped
egg production in 10 d.
Decreased photoperiods during the molting period
have been shown to hasten induction of molting and
reduce mortality during molting (Lanson and Smyth,
1955; Brake and Thaxton, 1982). Removal of water to
augment fasting has been recommended as part of many
molting techniques (Swanson and Bell, 1974b). Swanson
et al. (1978) reported that withholding water increased
egg size, which is undesirable in most cases. Palafox
(1976) did not find such an effect. At present, withholding
water is not recommended due to increased egg size con-
cerns and increased mortality (Brake and Carey, 1983).
MOLTING PERIOD AND RETURN
TO EGG PRODUCTION
Fasting Molts
Conventional fasting and photoperiod modification
techniques for molting result in a cessation of egg produc-
tion that varies according to the length of time that the
hens are fasted or photoperiod is reduced. Generally,
flocks fasted up to 10 d produce the first eggs within 5
wk and reach 50% production by 8 wk from initiation of
fasting (Wilson et al., 1969; Swanson and Bell, 1971,
1974b,c; Baker and Brake, 1981).
Low Ca Molts
Wakeling (1978) reported that birds that received a
0.09% Ca ration for 10 d returned to 50% production in
4 wk. Hens that received a 0.09% Ca diet for 14 d reached
50% production by the eighth week after molt initiation
(Douglas et al., 1972). Gilbert and Blair (1975) observed
that hens molted with a diet containing 0.05% Ca for 6
wk returned to 50% production by the eighth week after
initiation of treatment.
Low Na Molts
Whitehead and Shannon (1974) reported that pullets
fed a low Na ration for 7 wk molted and returned to 50%
production by 6 wk after the end of treatment. Nesbeth
et al. (1976a,b) observed that pullets brought to a complete
cessation of production by 23 d on a low Na diet returned
to production 17 d after the end of treatment. Hens that
did experience a decrease to below 10% production on a
low Na diet were observed to return to 50% production
within 2 wk of the end of treatment (Ross and Herrick,
1981).
High Zn Molts
Hens that received a Zn molting ration for 7 d resumed
production 1 wk after returning to a normal diet. Hens
973
that received a Zn ration for 12 d resumed production 25
d after removal of excess Zn from the diet (Scott and
Creger, 1976). Shippee et al. (1979) found that hens molted
with a Zn ration returned to production 2 wk from the
end of treatment and reached 50% production by 5 wk
from initiation of the molt. Arrington et al. (1967) found
that of the hens molted by use of a 28-d high I diet, 85%
returned to production within 10 d of removal of excess
I from the feed. The remaining 15% of the flock did not
resume production.
POSTMOLTING PERFORMANCE
Association of Performance
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with Reproductive Involution
Baker et al. (1981a,b) postulated that maximum involu-
tion of the reproductive organs is essential for optimum
postmolt performance. Variability of BW loss, and there-
fore differing degrees of reproductive involution, was
suggested as an explanation for the conflicting results
obtained by investigators using similar molting proce-
dures. Loss of primary flight feathers also has been used
as an indicator of the completeness of reproductive invo-
lution (Kreuger et al., 1977). However, some techniques
such as molting by high dietary I may not cause any loss
of feathers (Arrington et al., 1967).
Hansen (1960) and Len et al. (1964) reported an associa-
tion between improvements in both postmolting egg
quality and production and the length of the rest period.
The degree of improvement in postmolting performance
was associated with an increase in the number of days
during which no eggs were produced. Baker (1981) and
Baker et al. (1981a,b) found that improvements in post-
molting performance were associated with increasing BW
loss of hens up to 31% of original BW. Hens that lost the
greatest amount of weight exhibited the greatest improve-
ments in eggshell quality and highest egg production
after molting.
Reduction of BW beyond 35% may exert detrimental
effects on life of flock egg production. Although postmolt-
ing egg production continued to improve with increasing
BW loss beyond 30%, the recovery period, during which
no eggs were produced, was prolonged such that overall
egg numbers were reduced for the life of the flock (Carter
and Ward, 1981).
Livability
Livability in layer flocks is defined as the number of
hens remaining in the layer house and is expressed as a
percentage of the number of hens originally placed in the
house. Swanson and Bell (1974c) reported that the average
livability for 63 flocks molted by 10 d of fasting was 98.7%
during the feed withdrawal period. Wilson et al. (1969)
found that livability of fasted hens did not differ from
that of nonfasted hens. Improved livability of fasted hens
relative to nonfasted hens has been associated with re-
duced photoperiod during the fasting period (Brake and
974
Thaxton, 1982). Fasted hens that receive a photoperiod
of less than 12 h during the molting period exhibite im-
proved livability as compared to fasted or nonfasted hens
receiving a photoperiod of more than 12 h. Flocks molted
by way of low Ca rations have higher mortality as com-
pared to nonmolted controls (Douglas et al., 1972; Wake-
ling, 1978). Low Na molting rations were observed to
have no effect on livability as compared to nonmolted
controls (Whitehead and Shannon, 1974; Wakeling, 1978).
High Zn molting rations had no effect on livability as
compared to nonmolted controls (Shippee et al., 1979;
Palafox and Ho-A, 1980). Arrington et al. (1967) reported
that high I molting did not affect livability of molted hens
as compared to nonmolted controls.
Feed Consumption
Wakeling (1978) observed no differences in postmolt-
ing feed consumption among flocks molted by low Ca,
low Ca and Na, or low Na and controls. Hens molted by
low dietary Ca have been reported to exhibit reduced feed
efficiency compared to hens molted by fasting (Martin et
al., 1973). Little data is available in the literature concern-
ing the effects of high dietary Zn induced molts on feed
consumption and efficiency. For two separate studies, Zn
induced molting did not affect postmolting feed con-
sumption (Shippee et al., 1979; Palafox and Ho-A, 1980).
By definition, if feed consumption is not affected and egg
production increased then feed efficiency is improved. In
these studies, the performance of hens molted by high
dietary Zn satisfied this definition.
PHYSIOLOGICAL EFFECTS OF MOLTING
An extensive, but nevertheless incomplete, body of lit-
erature documents the physiological phenomena associ-
ated with molting of feathers and changes in reproductive
function. Changing day length is characterized as the
primary environmental agent responsible for inducing
molt in many avian species (Rowan, 1926). Photoperiodic
changes induce and entrain circadian and circannual
rhythms. These rhythms, manifested by the neural and
endocrine systems, initiate physiological processes that
produce migratory fattening and restlessness, changes in
reproductive function, and molting of feathers (Follett,
1973). In species exhibiting an annual molt, increasing day
length promotes gonadal development and reproductive
function. Decreasing day length initiates gonadal regres-
sion and allows molting to occur (Meier and Farner, 1964;
Pittendrigh and Minis, 1964; Wolfson, 1966; Follett and
Sharp, 1969; Meier and MacGregor, 1969; King, 1970).
Other factors, such as temperature, are thought to modify
the effects of photoperiod (Follett, 1973). Considerable
differences exist in the timing of feather replacement and
reproductive function among species, and there is no
general agreement as to the physiological entities respon-
sible for the initiation of feather replacement and involu-
tion of the reproductive tract.
BERRY
Natural Molting
In the jungle fowl (Gallus gallus), the wild progenitor
of the laying hen, establishment of broodiness appears to
be the primary initiating factor. With the onset of broodi-
ness, the jungle fowl hen experiences involution of the
reproductive organs, a halt to egg laying, and, eventually,
the shedding and replacement of feathers (Sherry et al.,
1980).
The broody jungle fowl hen exhibits a voluntary reduc-
tion in food intake, which has been referred to as sponta-
neous anorexia (Mrosovsky and Sherry, 1980). During
incubation of her eggs, the jungle fowl hen consumes
very little food and water even if it is placed near the
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nest. During this time, the hen loses approximately 20%
of her BW. Approximately half the BW loss is due to
involution of the reproductive organs (Sherry et al., 1980).
After the chicks have hatched, the hen gradually resumes
feeding and feather replacement commences.
Spontaneous anorexia associated with incubation and
brooding is a common adaptation of birds living in envi-
ronments where rearing of offspring is incompatible with
feeding. Other birds which nest in environments that
require a high degree of parental attention during incuba-
tion and brooding experience some degree of anorexia.
A number of grouse, quail, and pheasant species reduce
food consumption during incubation (Breitenbach and
Meyer, 1959; Anthony, 1970; Redfield, 1973). Geese and
ducks also reduce their feed consumption and lose BW
during incubation (Korschgen, 1977; Ankney and MacIn-
nes, 1978).
The most extreme example of spontaneous anorexia in
birds is that of the male emperor penguin (Aptenodytes
forsteri). The male emperor penguin is responsible for
incubating the sole egg laid by their mate during the
Antarctic winter. During this time, it is difficult for the
male to move for feeding or other purposes. Thus, these
birds will fast for about 120 d of incubation and brooding
during which time they lose 40% of their BW (Le Maho,
1977). From these examples, it is clear that many bird
species, including the chicken, survival with little or no
food for relatively long periods of time is a normal feature
of their physiology.
Endocrine Effects of Molting
Prolactin. There is a considerable body of evidence
implicating prolactin in the development of broodiness
and the subsequent collapse of reproductive function.
Prolactin concentration in the blood rises as egg laying
proceeds. Eventually, the high level of prolactin reduces
hypothalamic release of gonadotropin-releasing hormone
(GnRH) and release of luteinizing hormone (LH) from
the pituitary (Tabibzadeh et al., 1995). It appears that
prolactin may also directly inhibit ovarian steroidogene-
sis (Rozenboim et al., 1993). Taken together, these actions
of prolactin would lead to involution of the ovary with
reduced ovarian steroidogenesis and regression of the
oviduct.
 SYMPOSIUM: INDUCED MOLTING IN LAYERS
In species that exhibit broodiness, such as the jungle
fowl and the domestic turkey hen, the endocrine changes
that occur at molt induction begin with rising prolactin
levels. Eventually this inhibits the hypothalamic-pitu-
itary-gonadal (HPG) axis. However, in the commercial
laying hen induced to molt by fasting, plasma prolactin
concentration increases only slightly or not at all (Scanes
et al., 1979; Verheyen et al., 1983).
Corticosterone. The first significant endocrine change
observed in fasted hens is that corticosterone levels in-
crease as the hypothalamic-pituitary-adrenal (HPA) axis
is activated by the need to mobilize body energy stores.
Juhn and Harris (1956) observed that exogenous deoxy-
corticosterone was as effective as progesterone in induc-
ing molting. Perek and Eckstein (1959) inferred adrenal
activation in molting from the finding that adrenal
ascorbic acid is depleted in molting hens. Reports by
others (Beuving and Vonder, 1978; Brake et al., 1979)
indicate very little or no effect on adrenal activity of hens
molted by fasting. Also, circulating levels of adrenal corti-
cal hormones decreased during molting when photope-
riod is reduced (Gildersleeve et al., 1979). Etches et al.
(1983) reported that corticosterone is elevated in hens
during molting induced by fasting; however, the mea-
sured amount of corticosterone is lower than the amount
of exogenously administered corticosterone required to
induce ovarian regression.
The degree of the increase in corticosterone depends
on the method used to induce molt. Methods such as
fasting, which induce a rapid and complete molt, are
associated with larger increases in corticosterone levels,
whereas methods that take longer to induce molt result
in a smaller increase in corticosterone levels (Etches,
1983). The corticosterone increase is transitory and de-
creases toward prefast levels as fasting continues. Upon
refeeding, corticosterone levels again increase.
In the fasting hen, increased corticosterone levels are
associated with inhibition of the HPG axis (Williams et
al., 1985). Luteinizing hormone, and presumably follicle-
stimulating hormone, levels decline as corticosterone lev-
els increase at the beginning of the fast. A possible mecha-
nism for this is that corticosterone makes the pituitary
refractory to GnRH. In the laying hen, corticosterone lev-
els exist in an inverse relationship with LH (Wilson and
Cunningham, 1980). Diurnal cyclicity of corticosterone
secretion declines as hens become sexually mature, dur-
ing which time LH levels increase to maximum. However,
the interaction of the HPA axis and the HPG axis is com-
plex. In the hen, corticosterone may either augment or
inhibit reproductive function depending on the season,
environmental circumstances, and availability of feed
(Carsia and Harvey, 2000).
Ovarian Steroids. Loss of ovarian steroid support re-
sults in regression of the oviduct (Himeno and Tanabe,
1957; Tanabe et al., 1957; Tanabe and Katsuragi, 1962).
Ovarian regression begins soon after the start of fasting
with declines in estradiol and progesterone (Tanabe et
al., 1981; Etches et al., 1984). Exogenous treatment of do-
mestic fowl with antigonadotropic compounds such as
975
progesterone or prolactin will induce molting of feathers
and reproductive involution (Adams, 1955; Shaffner,
1955; Juhn and Harris, 1956, 1958; Tanabe and Katsuragi,
1962). Stake et al. (1979) used the antiestrogenic com-
pound, tamoxifen, to produce ovarian regression. Pino
and Hudson (1953) and Tanabe et al. (1957) induced molt-
ing with the antigonadotropic compound enheptin. Meth-
allibure, another antigonadotropic compound, has been
reported to have effects similar to enheptin (Ishigaki et
al., 1971). Castration of mature domestic fowl results in
a continuous molt, which is reversed by administration
of exogenous sex hormones (Assenmacher, 1958; Voitkev-
ich, 1966). These findings suggest that sex steroids inhibit
molting in domestic fowl. Hypophysectomized birds molt
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feathers intensely 2 wk after the pituitary is removed and
remain in a state of perpetual feather molt and gonadal
regression (Assenmacher, 1958).
In any case, exogenous administration of androgens,
estrogens, and synthetic estrogenic compounds inhibit
molting of domestic fowl (Takewaki and Mon, 1944; Ko-
bayashi, 1954; Vaugien, 1955; Wagner and Muller, 1963)
as does gonadotropin treatment in hypophysectomized
birds (Assenmacher, 1958). As LH and follicle stimulating
hormone levels rise following refeeding and photostimu-
lation, ovarian follicles enter into the maturational hierar-
chy and begin producing estradiol and progesterone.
Exposure to short day length and reproductive tract
involution abolishes photorefractoriness in seasonal
breeding birds (Etches, 1996). Induced molting serves this
purpose in the domestic hen. Upon returning feed to the
hens and resuming their exposure to long day lengths
following induction of molt, hypothalamic GnRH release
is resumed, and pituitary responsiveness is restored. This
results in increasing LH concentration, which stimulates
ovarian development and ovarian steroid production. Es-
trogen and progesterone levels increase. Corticosterone
levels decline, but may show a transient increase as egg
production resumes.
Thyroid Hormones. Many of the investigations into
the endocrine mechanisms of molting have focused on
feather molting induction. Early research suggested that
the thyroid gland controlled molting of feathers. Zava-
dovsky (1925a,b) reported that feeding mammalian thy-
roid glands to chickens results in feather loss and caused
male plumage to resemble female plumage. Following
treatment with whole, minced thyroid glands, birds ex-
hibit a refractory period during which further treatment
does not initiate a new cycle of feather loss. Similar results
were reported for an experiment utilizing White Leghorn
hens and cocks (Cole and Hutt, 1928). Van der Meulen
(1939) proposed that thyroid activity was primarily re-
sponsible for feather loss and replacement. However, he
also observed that thyroid treatment did not affect normal
reproductive cycles. From this, it was concluded that
feather loss and reproductive function are under sepa-
rate control.
Perek and Sulman (1945) observed an increase in the
basal metabolic rate concomitant with molting in Single
Comb White Leghorn hens. Thiouracil, an antithyroid
976
compound, was found to retard molting (Glazener and
Jull, 1946). However, Himeno and Tanabe (1957) reported
that feeding thiouracil could precipitate molting. Further-
more, Perek and Sulman (1945) observed that although
thiouracil treatment lowers the basal metabolic rate, it
does not prevent naturally occurring molting of Leghorn
hens. However, I uptake studies indicated that if hens
are maintained at elevated temperatures during periods
of feather loss then no increase in thyroid activity occurs
(Tanabe and Katsuragi, 1962). Also, earlier work by Ta-
nabe et al. (1958) indicated no increase in pituitary thyro-
tropin release in molting hens. In contrast to this observa-
tion, histological examinations revealed that thyrotropic
B cells predominate in the pituitary at the time of molting
(Perek et al., 1957). In addition, Brake et al. (1979) ob-
served an increased thyroxin (T4) level during fasting.
Refeeding induced a sharp decrease of T4 and an increase
in plasma triiodothyronine (T3) concentration.
Coincidence of feather molting and reproductive invo-
lution in the hen, in concert with evidence of thyroid
participation in molting, has led to considerable interest
in possible interactions of ovarian steroids with thyroid
hormones. Shaffner (1954, 1955) and Adams (1955) ob-
served that although progesterone administered to hens
precipitates molting of feathers and cessation of repro-
duction function, the simultaneous administration of thy-
roprotein acts synergistically to enhance the antigonado-
tropic effects of progesterone and to induce molting. Fur-
ther work by Brake et al. (1979) indicated that the T4
increase during fasting coincides with active regression
of the ovary. At the point of complete ovarian regression,
T4 decreases sharply with a concomitant increase in T3.
Vitamin D Endocrine System. The hormonally active
form of vitamin D, 1,25 dihydroxycholecalciferol
(1,25(OH)2D3), is of central importance to Ca transport
mechanisms in the shell gland and gut. In combination
with estrogen, 1,25(OH)2D3 is responsible for the Ca-bind-
ing protein calbindin in the intestine and shell gland.
Plasma concentrations of 1,25(OH)2D3 decrease as hens
age (Abe et al., 1982). Upon the return to egg production
following molting, plasma levels of 1,25(OH)2D3 are in-
creased to levels similar to that of young hens (Abe et
al., 1982). Receptors for 1,25(OH)2D3 in the shell gland
are also increased following an induced molt (Berry and
Brake, 1986). In addition, cellular proliferation in the shell
gland is correlated with increased tissue receptivity to
1,25(OH)2D3 and increases Ca-binding protein content
(Heryanto et al., 1997a). Therefore, induced molting in-
creases the level of the hormonal form of vitamin D and
tissue sensitivity to the hormone.
Effect of Molting on Organ Systems
Ovary and Oviduct. Loss of gonadotropin support
during fasting causes involution of the ovary. Follicles in
the maturational hierarchy become atretic and the yolk
material is resorbed. Ovary weight declines as follicles
become atretic. Reduction in ovary weight is initially de-
BERRY
pendent on the duration of fasting and the rate of BW
loss. Beyond 25% BW loss, the ovary is fully regressed.
Involution of the oviduct follows the loss of ovarian
steroidal support. Regression of the oviduct is a true re-
modeling of the tissue rather than a decline in the size
of cells or shrinkage of the tissue. Apoptosis removes
cells of the glandular epithelium during regression (Hery-
anto et al., 1997b). Remodeling even extends to the con-
nective tissue of the oviduct as evidenced by increased
levels of collagenase activity during involution.
Upon resumption of sex steroid production by the
ovary during recovery from the molt, the oviduct recru-
desces. Relatively little effort has been expended to study
the physiological basis for the apparent rejuvenation of
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egg production and eggshell quality that occurs following
molting. Brake and Thaxton (1979) postulated that “...the
effects of this rejuvenation may be manifested in many
physiological parameters; however, discrete changes at
the cellular level must be the driving force. This rejuvena-
tion may be mediated by an increasing tissue sensitivity
or efficiency. Reorganization of metabolic processes, re-
moval of inhibiting substances at the cellular level, or
reversal of the normal, but as yet undefined, aging pro-
cesses of the laying hen could be involved.”
Improved shell gland function following induced molt-
ing may be due to remodeling at the cellular level. Cellular
proliferation in the oviduct replaces cells lost during the
regression, as evidenced by increased staining of the pro-
liferating cells for proliferating cell nuclear antigen, a
marker of cell proliferation (Heryanto et al., 1997a).
Remodeling of shell gland tissue may also be responsi-
ble for removing substances that interfere with shell gland
function. The lipid content of the shell gland increases as
hens age. Induced molting reduces shell gland lipid and
alters the balance of lipid types (Baker et al., 1980; Baker,
1981). Evidence for changes at the cellular level in the
oviduct were found in reports from several researchers.
Baker (1981) observed that the uterine glandular epithe-
lium, which is the site of eggshell Ca transport and deposi-
tion, contain quantities of intracellular lipid visibly detect-
able by histological staining. Roland et al. (1977) reported
that hens laying shell-less eggs have significantly higher
uterine lipid levels compared to hens producing normal
eggshells. Further reports by Baker et al. (1981a) indicated
that induced molting halts the incidence of shell-less eggs
and removes lipid accumulation in the uterus. Baker et
al. (1980), Brake et al. (1981a), and Brake and McDaniel
(1981) have reported that optimum postmolting perfor-
mance was achieved when BW loss was greater than 25%.
They reported that uterine lipid was not lost until BW
decreased to that point, which coincided with maximum
oviductal regression.
As noted previously, molting increases tissue receptiv-
ity to 1,25(OH) dihydroxyvitamin D3 (Berry and Brake,
1987). The location of the cytosolic receptors for 1,25(OH)
dihydroxyvitamin D3 in shell gland glandular cells coin-
cided with the reported location of Ca binding protein.
Intestine. Duodenal weight decreases during fasting
and returns to original size upon refeeding. Following
 SYMPOSIUM: INDUCED MOLTING IN LAYERS
the molt, intestinal Ca binding protein concentration in-
creases (Berry and Brake, 1991) compared to unmolted
hens. Intestinal uptake of Ca also improves following
molt (Al-Batshan et al., 1994).
Immune and Hematopoietic Systems. Significant
changes in the immune and hematopoietic systems have
been noted in molting hens. Reproductive quiescence in
the hen is accompanied by recrudescence and lympho-
cytic repopulation of the thymus (Brake et al., 1981b).
Thymic recrudescence and lymphocytic repopulation has
been observed during the reproductive quiescent periods
of other seasonal breeding animals such as deer. It is not
known whether this serves an adaptive function or is
merely the result of removal of gonadal steroid inhibition
of thymic function. Peripheral lymphocyte numbers de-
crease and lymphocyte populations change during the
fasting period of the molt (Alodan and Mashaly, 1999).
The proportion of CT 4 lymphocytes decrease while the
proportion of CT 8 lymphocytes increase (Holt, 1992),
and a relative heterophilia has been consistently noted in
molting hens (Holt, 1992). However, heterophil chemo-
taxis, phagocytosis, and oxidative burst are inhibited.
A decrease in the number of B cells has been observed in
hens during fasting-induced molts (Holt, 1992). However,
antibody levels do not appear to be changed. These
changes are similar to those observed in birds placed in
stressful conditions and are also similar to cellular
changes observed during inhibition of immune function
in mammals following trauma. This would appear to
reduce the birds’ ability to respond to new antigenic chal-
lenges. In fact, conventional wisdom is that hens should
not be vaccinated during molt. Reduced immune function
during molt may contribute to the reported increase in
Salmonella incidence in hens during induced molt (Porter
and Holt, 1993; Holt et al., 1994).
Packed red blood cell volume (hematocrit) increases
during the period of reproductive involution and de-
creases again as the reproductive tract regenerates. This
is due to removal of estrogen inhibition of red blood
cells (RBC) formation, reduced vascular volume due to
involution of the ovary and oviduct, and presumably to
decrease fluid intake leading to hemoconcentration. Molts
induced by high levels of dietary Zn are an exception.
Packed cell volume does not increase or may decrease as
RBC are removed from circulation (Berry et al., 1987).
Numbers of RBC decline as ovarian function returns
due to estrogen inhibition of RBC production, expansion
of vascular volume, and hemodilution. Reticulocytosis is
observed during molt recovery. During recovery from
Zn molts, the reticulocytosis is extreme and packed cell
volume may show a transient increase at the time when
packed cell volume is actually declining in fasting molted
hens (Berry et al., 1987).
CONCLUSIONS
Induced molt produces discreet changes at the endo-
crine and cellular levels that improve egg production and
eggshell quality. Increased circulating concentrations of
977
1,25(OH)2D3, increased tissue 1,25(OH)2D3 receptor con-
centrations, and increased egg shell gland and intestinal
calbindin-D28k concentrations are examples of these en-
docrine and cellular changes.
Molting is also associated with thymic recrudescence
and lymphocytic repopulation that may improve immune
function in the hen. However, during the induced molt,
there are also transient reductions in the number of spe-
cific lymphocyte classes that may cause an increased sus-
ceptibility to infection. Reduced mortality during and
after induced molting suggests that molting does not im-
pair immune function with respect to avian pathogens.
However, increased susceptibility to colonization by or-
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ganisms such as foodborne Salmonella, which are not
necessarily avian pathogens, are nevertheless of concern
from a food safety perspective.
Animal welfare activists have objected to induced molt-
ing on the grounds that fasting to induce molting is cruel.
However, the degree of fasting imposed to induce molt
must be viewed in the context of the physiological adapta-
tions of the hen. Most birds are adapted to exploiting
food sources that are unevenly distributed both time and
space, forcing the birds to survive without food for in-
definite periods. Many species are also required to fast
for extended periods by the conditions of migration or
incubation. These birds have a number of adaptations
including the ability to build fat reserves. Besides reliance
on catabolism of abdominal and other fat depots, the
jungle fowl and the commercial hen have the ability to
consume resources made available by involution of the
ovary and oviduct which spares other fat and protein
reserves. It is clear that many avian species, such as the
jungle fowl, are well adapted to survive prolonged fast-
ing. Likewise, it would appear that the commercial layer
is tolerant of fasting as evidenced by reduced mortality
during the fasting phase of induced molts.
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