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Early Feeding Immune Development Neonatal Poultry': System
Early Feeding Immune Development Neonatal Poultry': System
Early Feeding Immune Development Neonatal Poultry': System
1 Presented at the 1998 Poultry Science Association Informal Poultry Nutrition Symposium:
"Impact of Early Nutrition on Poultry."
2 To whom correspondence should be addressed
426 EARLY FEEDING AND IMMUNITY
options, for example during shipment of to respond to a vaccine with the development
birds over long distances. of immune memory. Studies will be discussed
Producers are well aware that post-hatch with reference to these three postulates.
holding and bird processing are hard on the
hatchlings and take steps to minimize their AND METHODS
MATERIALS
effects. Many hatcheries attempt either to
place the birds within hours of removal from In these studies, the effect of early feeding
the hatcher or to hold them for a sufficient on the immune system of broiler chicks [3]
period of time to mature and recover from was evaluated by measuring the weights of
processing. This rest and recovery from pro- immune organs and levels of serum and biliary
cessing has an additional benefit. Just before immunoglobulin A (I@) [4],and by evalua-
the hatching process begins, the bird inter- tion of cell proliferation and immunoglobulin
nalizes what is left of the yolk sac, which has isotype expression by lymphocytes. The nutri-
nourished it during incubation. The residual ent source fed to these neonatal chicks was
yolk protein is the source of antibodies from a hydrated nutritional supplement (Oasis
the hen [l]. To be effective, maternal anti- hatchling supplement) consisting of 70%
bodies must not only move from the residual water, 10% protein, 20% carbohydrate, and
yolk into the bloodstream but must also diffuse less than 1% fat [q.This hydrated nutritional
to sites of vulnerability - in particular to the supplement (HNS) was replenished daily and
mucosal surfaces where organisms are most was fed to the treated birds ad libitum on the
likely to enter the body. Thus a delay in place-
day of hatch (Day 0) and the two subsequent
ment can leave the hatchlings better able to
respond to the environment once they are days (Days 1and 2). Control birds were fasted
placed. and given no water over the same period. Be-
The attendant problem is that birds are ginning on Day 3, all birds were given water
generally not fed in the hatchery - even when and fed an identical corn-soy starter diet for-
held overnight - nor are they fed during trans- mulated to meet or exceed National Research
port. Producers may feel that feeding is not Council recommendations for starter feed [6].
essential during this period because conven- Effects on performance, organ weights, im-
tional wisdom says that the bird can survive on munoglobulin expression, and levels of serum
its residual yolk [2]. This is a valid statement and biliary IgA were measured in four birds
but does not completely represent the modern per treatment on Days 0, 1,2, 3,6, 7,8,9, 10,
chick or poult. While the survivalof a hatchling 13, 14, and 21. Tissue sections of small intes-
may indeed depend, in the absence of other tine, ileocecal junction, bursa, and thymus
feed, upon its use of residual yolk as a nutrient were prepared and stained using hematoxylin
source, the research described in this report and eosin for purposes of morphometry. To
indicates that this is not the optimum use for evaluate microscopic structure, villus length,
residual yolk. In addition, data indicate that mid-villus width, crypt depth, and bursa folli-
the development of the immune system in cle area were determined. Hematoxylin and
particular appears to respond to early feeding. eosin staining and immunocytochemical
Three mechanisms are proposed to ac- methods for bromodeoxyuridine and IgA
count for the dramatic effect of oral nutrition
have previously been described and those for
on the hatchling immune system. First, early
nutrition may provide limiting substrates; immunoglobulinsM (IgM) and G (IgG) differ
second, feeding may affect endogenous levels only slightly [A.
of hormones or other immunomodulators;and In a separate study, birds were fasted or
third, the presence of antigen in the gastro- were fed the hydrated nutritional supplement
intestinal system may be necessary to trigger for the day of hatch and the day after. All birds
full differentiation of the primary immune were then allowed to consume the same corn-
cells, particularly the B lymphocytes. Com- soy starter diet ad libitum. Buds were chal-
plete differentiationof these cells is critical for lenged orally with a 1OOx dose of a commercial
the eventual development of secondary im- coccidiosis vaccine [8] on Day 14, i.e. after
mune structures such as germinal centers or 12 days on ad libitum feed. Birds and feed
cecal tonsils, along with the associated ability were weighed on Days 7,14,and 20.
Symposium
DIBNER et al. 427
FIGURE 1. Bursa of Fabriciuson the day of hatch stained for: A) immunoglobulin M (IgM); B) immunoglobulin
G (IgG); and C) immunoglobulin A (IgA). The B lymphocytes of the neonatal chick can synthesize only IgM. All
of the IgG is found in blood vessels and represents maternal antibody. There is no indication of IgA, of either
maternal or chick origin.
-
h
E
cn
.-
125
cn
Sa ’
- - Fasted
*Fed HNS
0.75
Y I
8
m a
0.5 8
-1GURE 2. Bursa weight as a function of age in fasted birds or birds fed a hydrated nutritional supplement
:HNS)for post-hatch Days 0, 1, and 2. Bursa weights were significantly heavier (P c .OOOl) for the 21-day period
or the birds fed HNS (SEM= .05; *Means significantly different, P < .05).
Symposium
DIBNER et al. 429
0.3
0.25
F
E0
-Es 0.2
.$ 0.15
sm
m
$ 0.1
0.05
0
0 1 2 3 8 15 21
Day of Age
FIGURE 3. Bursa weight as a percentage of body weight in fasted birds or birds fed a hydrated nutritional
supplement (HNS) for post-hatch Days 0, 1, and 2. Bursa weight as a percentage of body weight
was sionificantlv heavier I P e .003)for the birds fed HNS from Day 3 through Day 21 (SEM = .03;*Means
significantly diffkrent, P e .05). ’
bursa weight may simply be the rise in surplus even in birds that are fed immediately
glucocorticoids associated with fasting [24]. after hatching.
Glucocorticoids have been reported to be as- The development of secondary immune
sociated with involution of primary lymphoid tissues such as the spleen, cecal tonsils, and
organs, even in young poultry [25]. Harderian gland is clearly dependent first on
Low substrate availability or low oral in- the bursa and thymus [31]. However, in con-
take of antigen could also cause the results trast to these primary immune organs, a criti-
shown in Figures 2 and 3. Although the im- cal influence on the development of secondary
mune response does not appear to be limited tissues appears to be antigen exposure [32,33].
by substrate availability later in life, nutrient The cecal tonsils are a very good example of
requirements for the development of the im- this, with germ-free animals demonstrating
mune system have not been similarly exam- small cecal tonsils devoid of germinal centers
ined. During the frrst week of life, substrate [34]. This effect lasts through about 4 wk, after
availabilitymay well be limiting. The growth of which a few small germinal centers can be seen
certain systems, particularly the gastro- [35]. Eventually, some of the animals may
intestinal, cardiovascular, and respiratory, is exhibit low levels of serum IgG, but there are
critical to the achievement of the genetic
no reports of the presence of IgA. Germ-free
potential of the bird for growth. Modern
animals can also be deficient in isotype differ-
production systems and genetic selection for
performance in poultry may have diminished entiation and at the extreme may be limited
the bird’s immune responsiveness and may to an IgM response with very low antigen-
also have influenced partitioning of nutrients binding diversity [%, 371. It is interesting that
in the neonate [26, 271. The growth rate of this difference between conventional and
the gastrointestinal system over the first week germ-free animals exists, because the feed
of life has been estimated at three to five itself, even if sterile, should provide some
times that of the rest of the body [28, 291. In antigen. It may be that antigens from living
addition, nutrient transport systems, gut sur- microorganismsare required for this function.
face area, and digestive enzyme levels have In any case, feeding on the day of hatch may
been found to be adequate but not in excess provide an early antigen stimulus and thus
of early need for nutrients [30]. These obser- facilitate rapid differentiation of the humoral
vations suggest that there may not be a nutrient response [38,39].
430 EARLY FEEDING AND IMMUNITY
Figure 4 shows sections of bursa from lymphocyte proliferation does not prove that
birds fasted or given the HNS for 3 days. The it results in an improvement in development of
sections are stained to detect proliferating immunocompetence. Other observations,
cells, whose nuclei appear dark. Note that in however, suggest that there may be long-term
the bursa from the fed bird, virtually every consequences of early feed and water depriva-
lymphocyte is stained. The bursal lymphocytes tion. Figure 5 shows the effect of early feeding
undergo an explosive proliferation in the on the appearance and levels of biliary IgA.
neonate, as long as substrate and antigen are This immunoglobulin is a part of the mucosal
present. Figures 3 and 4 illustrate, and it is immune system and is the last of the major
important to emphasize, that the contents of isotypes to appear. Thus, presence of IgA is a
the residual yolk cannot be substitutedfor oral sign that the humoral immune system is fully
intake. First, note that the animals in this study developed. As is clear from Figure 5, early
were not deutectomized and all had residual feeding was associated with a more rapid ap-
yolk available a5 a source of nutrients. Appar- pearance of biliary IgA and a generally higher
ently, there was not enough nutrient capacity level of the immunoglobulin over the entire
in the yolk to maintain lymphocyte prolifera- 21-day study.
tion at the same level as seen in birds receiving Another indicator of immune maturation
oral nutrition (Figure 4). This may be due to is the appearance of germinal centers [14].
the priority use of nutrients by the gut and These are local concentrations of lympho-
cardiovascular and respiratory systems but cytes in which T cells, B cells, and antigen-
may also be related to a lack of antigen from presenting cells form an organized structure
the gastrointestinal system. associated with the development of immune
memory to antigen, such as is required for a
D E V E L O P M E N T O F SECONDARY vaccine response. In the study reported here,
RESPONSES AND DISEASE RESIS- germinal centers in the cecal tonsils were used
TANCE to indicate full immune maturation and the
The demonstration that early feeding in- capability to mount an anamnestic response.
creases bursa weight and the amount of bursal Figure 6 shows the effects of early feeding on
r- 0.35-
0.3 -
025 -
4
0
-
z
j
w
0.2
0.15 -
O
0.05
0
0 2
---.
4 6
.
a
.
'
8 10 12
l
14 16 18 20
~
22
,
Age (Days)
FIGURE 5. Biliary immunoglobulin A (IgA) levels as a function of age in fasted birds or birds fed a hydrated
nutritional supplement (HNS) for post-hatch Days 0, 1, and 2. Data points represent a pooled sample from four
birdsfireatment/day, and no estimate of variability was obtained.
- -Fasted
+Fed HNS
FIGURE 6. Number of germinal centers in sections of cecal tonsil as a function of age in fasted birds or birds
fed a hydrated nutritional supplement (HNS) for post-hatch Days 0, 1, and 2. Number of germinal centers was
significantly greater (P<.OoOl) for the 2 l d a y period for the birds fed HNS (SEM=.9; *Means significantly
different, P < .05).
the appearance of germinal centers. It is clear disease resistance. These birds had not been
that there is an effect of early feeding and that immunized for coccidiosis and all effects
once these structures appear they undergo a shown were due to the presence or absence of
linear increase with age. nutrients on Days 0 and 1.As can be seen in
Finally, the effect of early feeding on dis- Figure 7, the fasted birds were not as heavy as
ease challenge was tested. In the study shown the early fed birds, even 18 days after all the
in Figure 7, a challenge using a commercial buds were consuming the same diet ad libitum.
coccidiosis vaccine [a] was used to evaluate In addition, there was a significant difference
JAPR
432 EARLY FEEDING AND IMMUNITY
12.0
0.75
m
5 0.5
Em
.-
iz
3 0.25
m
0
Body Weight Cumulative Feed to Gain
ZIGURE 7. Performance of birds fed or fasted over post-hatch Days 0 and 1 and then placed on a corn-soy diet
&libitum. (Treatmenteffects: bodyweightP<.0003, SEM=.Ol; cumulativefeedefficiencyP=.l4,SEM=.W;
=Means significantly different, P < .05).
in performance between fasted and fed birds the response. For example, a period of feed
following the coccidiosis challenge. Birds fed restriction in poultry has been reported to in-
the HNS on Days 0 and 1 retained the im- crease the cellular and humoral response to
proved performance associated with feeding sheep red blood cells [42]. Specific nutrients
even during a disease challenge. It should be affecting the immune response have also been
emphasized that this effect was not exclusive identified. As an example, dietary fatty acids
with respect to coccidiosis. The oral challenge have been reported to affect the levels and
was simply used as a model for a non-specific types of responses to an immune challenge
stress or disease challenge. The data suggest [43]. Dietary immunomodulators can amplify
that birds given the optimum nutrient formu- or diminish the magnitude of the reaction to
lation immediately after hatch are better able a challenge through their effects on other im-
to respond to the variety of physiological and mune cells [MI.
environmental challenges of the production Development of the avian immune system
facility.This was also observed in turkey poults has been widely studied, but little informa-
exposed to a challenge model for poult enter- tion has been published on the effects of early
itis and mortality syndrome [41]. feeding on its development. There are three
NUTRITION AND IMMUNITY ways in which early feeding could affect im-
The studies reported here are intended to mune development. First, nutrients provide
clarify the effect of early nutrition on the de- substrates for cell proliferation and differ-
velopment of the immune system. It is import- entiation; second, nutrients can be im-
ant to distinguish immune development from munomodulators themselves or can affect
the immune response. Numerous publications their endogenous synthesis; and third, oral
have covered the subject of nutrition as it intake provides many of the antigens that drive
affects the ability of the animal to respond to both the development of isotypes and the gen-
an immune challenge. Nutrition can affect the eration of immunoglobulin diversity in the
magnitude of the response and the nature of bursa [45,46,47l.
Symposium
DIENER et al. 433
CONCLUSIONS
AND APPLICATIONS
Development of the immune system is initiated during embryogenesis but is not complete
until weeks or months after hatch. This development may be limited by nutrient availability
in fasted hatchlings.
Early feeding was associated with larger bursa welghts and greater lymphocyte prolifera-
tion. Residual yolk did not provide the required level of nutrition to fully support immune
system maturation during the first two days after hatch.
Appearance of biliary IgA and germinal centers occurred earlier and in larger amounts in
birds given early nutrition, indicating a more rapid development of the capabilityto respond
to vaccine administration. Early feeding was associated with improved bird performance
following a disease challenge.
The chick or poult should be provided with an optimum nutrient formulation and a source
of water immediately after hatching. This initiates immune development and spares yolk
macromolecules such as yolk antibodies for passive immunity. Biochemically, residual yolk
lipids are ideal for Lipid transport, for cell membrane and immunomodulator synthesis, and
for development of the central nervous system and retina.
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