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Lipid and carbohydrate sources for elemental, enteral diets for neonates

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Lipid and carbohydrate sources for elemental, enteral diets for neonates
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Rice, Lori P., 1957-
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ix, 147 leaves : ill. ; 29 cm.

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Fats ( jstor )
Foals ( jstor )
Horses ( jstor )
Infants ( jstor )
Milk ( jstor )
Neonates ( jstor )
Piglets ( jstor )
Plasmas ( jstor )
Small intestine ( jstor )
Swine ( jstor )
Animal Science thesis Ph. D
Dissertations, Academic -- Animal Science -- UF
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bibliography ( marcgt )
non-fiction ( marcgt )

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Thesis:
Thesis (Ph. D.)--University of Florida, 1989.
Bibliography:
Includes bibliographical references (leaves 138-145).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Lori P. Rice.

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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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LIPID AND CARBOHYDRATE SOURCES FOR ELEMENTAL,
ENTERAL DIETS FOR NEONATES



By

LORI P. RICE


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY

















UNIVERSITY OF FLORIDA


1989




LIPID AND CARBOHYDRATE SOURCES FOR ELEMENTAL,
ENTERAL DIETS FOR NEONATES
By
LORI P. RICE
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1989


This dissertation is dedicated to my father,
Bernard Rudin,
who passed away in November, 1987. His humor,
wisdom, support, and encouragement, unselfishly
given to all who knew him, inspired great deeds.
He believed that I could accomplish anything I
set out to do. All things were possible.


ACKNOWLEDGMENTS
Sincere gratitude and appreciation are extended to the
chairman of my supervisory committee, Dr. E. A. Ott. He
always had patience, tolerance, and good spirits, even
during difficult times (such as trying to catheterize an
uncooperative foal at 3 a.m.). His guidance and knowledge
were essential to the completion of this project.
A special thanks is given to Dr. Peggy Borum, who
introduced me to the challenging field of neonatal
nutrition. Her dedication and high research standards were
an inspiration to do the very best job possible.
Support, encouragement, and advice were liberally given to
me by the other members on my advisory committee: Dr. E.
Johnson, Dr. D. Beede, and Dr. S. Lieb.
I am extremely grateful for the assistance of Fred
Buhl, who solved all my computer problems, Dr. Wilcox, who
very patiently helped with the statistical analyses, Mel
Tooker, and the crew at the Horse Research Center (C. Roden,
farm manager, Rose, Cindy, Jim, and Sharon). Maria
Fernandez and Sandy Duyck performed miracles restraining
fiesty foals and provided vital feedback. The piglet
projects benefitted from the experienced help of Janet
iii


Baltzell, Mike Sullivan, Sergio Quintana, and Robin Adkins.
My friends, Ty and Scott McClung, have been especially
helpful and supportive during my entire program.
Unwavering support, love, and encouragement were given
to me by my family. My husband, Brett, allowed himself to
be dragged 1000 miles to sweltering Florida and put up with
a student for a wife for longer than he'd like to remember.
My mother, Ellie Rudin, my grandmother, Ida Rudin, my sister
and her husband, Gail and Chuck Willis, were always there
when I needed them.
IV


TABLE OF CONTENTS
Page
ACKNOWLE DGMENTS iii
ABSTRACT viii
CHAPTER
I REVIEW OF THE LITERATURE 1
Introduction 1
Formulating Enteral Diets for Neonatal
Foals 2
Foals as Research Animals 3
Piglets as Models in Foal Research 5
Digestive Problems of Neonates 7
Infectious Causes of Diarrhea 7
Non-Infectious Causes of Diarrhea 8
Lipid Metabolism in Neonates 9
Fat in the Neonatal Diet 9
Fatty Acid Digestion and Absorption 11
Metabolism of Medium Chain and Long
Chain Fats by Neonates 13
Disaccharidase Activity in Neonatal Small
Intestine 15
Introduction 15
Brush Border Disaccharidases 19
Developmental Patterns of Enzyme Activity
in Neonates 20
Distribution of Disaccharidases 25
Sucrase 26
Lactase 27
Maltase 28
Determining Disaccharidase Activity in
Experimental Animals 3 0
Responses of the Neonate to Stress 31
Failure to Acquire Passive Immunity 31
Cold Stress 33
Compromised Neonates 3 6
Hematology and Blood Chemistry in Neonates.... 37
Hematology 37
Chemistry 39
Milk Composition and Intake 4 0
v


Page
Milk Production in Mares 41
Special Nutrient Requirements of
Neonates 4 3
II EVALUATION OF AN ELEMENTAL, ENTERAL DIET FOR
NEONATES USING PIGLETS AS A MODEL 45
Introduction 45
Materials and Methods 48
Animals 48
Surgery 50
Diets 52
Development of Feeding Regimen 56
Daily Care 57
Tissue Collection 58
Analyses 59
Additional Information 60
Statistical Analysis 61
Results 61
General Observations 61
Gross Observations at Necropsy 64
Weight Gain 65
Small Intestine Length 66
Blood Glucose 67
Small Intestine Weight 67
Disaccharidases in the Small Intestine... 67
Complete Blood Counts 73
Urinalysis 75
Discussion 75
Disaccharidases in Neonatal Pigs 76
Growth Rate in Neonatal Pigs 79
Development of the Small Intestine in
Neonatal Pigs 79
Blood Glucose 80
Hematology 81
Summary of Piglet Trial 81
III FOAL ORAL DISACCHARIDE TOLERANCE TEST 8 3
Introduction 83
Materials and Methods 87
Preliminary Study 87
Animals and Design 88
Analyses 89
Statistical Analyses 89
Results 93
General Observations 93
vi


Page
Fasting Blood Glucose 94
Glucose Absorption Time Curves 94
Treatment Curves Pooled Over All Days.... 95
Comparison of Day-Treatment Curves 100
Discussion 108
Fasting Blood Glucose in Neonatal Foals.. 108
Treatment Curves Pooled Over All Days.... 110
Day 1 Treatment Curves 112
Day 3 Treatment Curves 112
Day 5 Treatment Curves 114
IV CONCLUSIONS 118
APPENDICES
A PIGLET AND FOAL DATA 122
B DISACCHARIDASE ASSAY PROCEDURE 129
LITERATURE CITED 13 7
BIOGRAPHICAL SKETCH 145
vii


Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
LIPID AND CARBOHYDRATE SOURCES FOR ELEMENTAL,
ENTERAL DIETS FOR NEONATES
By
Lori P, Rice
December 1989
Chairman: Edgar A. Ott
Major Department: Animal Science
Colostrum-deprived piglets were fed either elemental,
enteral diets, containing hydrolyzed whey proteins and
malto-dextrins (Group E), or a conventional milk-based diet
(Group C). Two litters were euthanized at birth (BO) to
provide baseline values. Normal values were obtained from
two litters of sow-fed (SF) piglets.
Group E piglets were surgically fitted with gastric,
bladder, and umbilical artery catheters, and fed by bottle
or continuous infusion pump. These piglets experienced
diarrhea. Diet and flow rate modifications, made for each
litter, reduced but did not eliminate the diarrhea.
To investigate diaccharidase activity in the small
intestine (SI) of these piglets, tissue homogenates were
incubated with maltose, lactose or sucrose. Maltase
activity was low in the SI of BO piglets suggesting that the
malto-dextrin in the elemental diet was probably not well
viii


digested by Group E piglets. Undigested maltose reaching
the large intestine probably caused the diarrhea.
Maltase activity was detected in the SI of Group C and
SF piglets even though they had never ingested maltose.
Similarly, lactose activity was detected in the SI of Group
E piglets that had never ingested lactose. Sucrase activity
was very low in all of the piglets.
To investigate disaccharidase activity in neonatal
foals, an oral tolerance test was conducted on 13 foals.
Maltose, lactose, sucrose, and glucose solutions were
administered orally on d 1, 3, and 5 postpartum. Changes in
plasma glucose from venous blood were measured over time.
Large peaks in plasma glucose occurred shortly after dosing
with lactose and glucose on all days. This suggests that
glucose from both substrates was absorbed. No rise in
plasma glucose was detected after oral sucrose
administration, even on d 5. Maltose was not well digested
on d 1. However, some digestion may have occurred on d 3
and 5, as evidenced by small increases in plasma glucose
after dosing with the maltose solution.
The colostrum-deprived piglet can be used successfully
as an animal model for nutritional investigations. The
results of the piglet and foal trials suggest that maltose
and sucrose would not be suitable substrates for these
neonates in the first few days of life.
ix


CHAPTER I
REVIEW OF THE LITERATURE
Introduction
Advances in medicine have decreased greatly the
mortality rate of weak or sick neonatal humans and animals.
New techniques in neonatology, especially in the field of
respiratory therapy, have enabled veterinarians to save many
diseased or premature foals that otherwise would not have
survived. The value, whether monetary or sentimental, of
many horses today has resulted in a greater number of high
risk foals receiving intensive care. As a result of the
great strides made in saving these foals, a growing number
of veterinary hospitals are developing or expanding their
facilities for the care of neonatal foals.
Once conditions that immediately threaten the life of
the newborn are stabilized, the primary concern becomes
administration of nutrients in a form that can be readily
utilized. In many cases, the foal can not or should not
consume mare's milk. Some foals must be separated from
their dams while they are in an intensive care setting.
Sick or weak foals that can not stand to nurse must be hand-
fed an appropriate enteral or parenteral diet.
1


2
Many of the feeding programs used on neonatal foals were
originally developed for human infants in intensive care
nurseries. Unfortunately, due to the difficulties in
working with these neonates, many of the formulas and
supplements being fed have not been well tested on very
young or stressed foals (Koterba and Drummond, 1985).
Formulating Enteral Diets for Neonatal Foals
Before an appropriate diet for stressed foals can be
formulated, basic research is needed to provide an
understanding of their metabolism. Many questions about the
efficiency of utilization of different nutrients remain
unanswered.
Ensuring adequate caloric intake is of primary
importance. In the past, some foals were inadvertently
starved to death while being hand-fed because their energy
requirements were underestimated (Koterba and Drummond,
1985). There is disagreement in the literature about the
amount and type of enteral diet that will supply adequate
nutrients during the critical neonatal period (Naylor and
Bell, 1985) .
Stressed or sick foals often have poorly functioning
digestive systems and may not absorb nutrients or antibodies
well. Some of these foals may even need parenteral feeding
to increase the supply of nutrients available to the cells.
Young foals are especially susceptible to infection when


3
they do not get an adequate supply of colostral antibodies
from their mothers, or when they are premature or dysmature
at birth (Rossdale, 1987). It has been shown in horses and
in other species that the nutritional needs of premature and
sick neonates can be different from that of term, healthy
newborns. While their need for nutrients remains the same
or even higher than that for healthy foals, their appetites
may be poor (Koterba and Drummond, 1985). Therefore, the
conclusions drawn from studies done on normal foals would
not necessarily be applicable to the weak or sick patients
treated in veterinary clinics.
Foals as Research Animals
Many problems present themselves when trying to design
experiments involving a relatively large number of research
foals. The expense of buying and feeding broodmares is
great, and there is no assurance that each mare will produce
a foal every year. Horses have a long gestation and tend to
foal during the same time of year. This makes it difficult
to spread foaling out over a long enough period of time to
facilitate handling many foals.
The labor required for round-the-clock care of even a
single foal is extensive. Even very young foals are
relatively large and require more elaborate facilities than
do small laboratory animals. Also, large radioactive
carcasses resulting from studies involving labeled compounds


4
would be difficult to dispose of. These concerns tend to
limit the number of foals that can be successfully studied
each year. These problems can be overcome, in part, by
using an appropriate animal model for preliminary
investigations.
Using animals as experimental models for human studies
is common practice. Ethical considerations have led to
extensive use of animal models for research concerning human
infants. Although the rat is frequently used as a model for
adults, rat pups are not similar to infants or foals.
Newborn rats have intestines that are fragile and difficult
to work with, and the rats' stage of development and rate of
growth at birth are very different from newborn humans,
pigs, horses, and ruminants (Widdowson, 1984). Calves,
while similar to foals, would not offer many advantages as a
model for foals.
Piglets have been used successfully as models for
premature infants. It has been suggested that they are
closer to human newborns than any other animal (Glauser,
1966). One of the objectives of this research is to
determine the appropriateness of using piglets as models
for the neonatal foal.


5
Piglets as Models in Foal Research
One of the major limitations of any animal model is, of
course, that the model is not going to behave exactly like
the real subject. Therefore the model chosen should always
come as close as possible to resembling the animal of
interest. There are no references in the literature to
previous work comparing piglets and foals, but information
on each shows that these neonates share many common traits.
The pig and the foal both tend to nurse freguently during
the day for short periods of time (Pegorier et al., 1983).
They have similar digestive functions at this age, both
being adapted to a milk diet. Also, both species depend
heavily on the passive immunity provided by the mother's
colostrum, which must be ingested shortly after birth.
It has been shown by many workers that foals and
piglets who do not acquire colostral antibodies are at
severe risk from many diseases including pneumonia,
septacemia, arthritis, and infectious diarrhea (Rossdale and
Leadon, 1975; Naylor and Bell, 1985; Rossdale, 1985; Koterba
et al., 1984; Mouwen, 1971; Bergeland and Henry, 1982). The
colostrum from both sows and mares contains predominantly
IgG (Pegorier et al., 1983; Naylor, 1979); however, IgA is
present in greatest quantities in non-colostral mare's milk.
Good management and sanitary conditions can help save
hypoglobulinemic neonates, but the risk of infection is
still high.


6
It has been suggested that mare's milk is closer to
that of primates than any other animal, being high in
lactose and low in fat and protein (Weaver, 1986). If
mare's milk is similar to human milk, and piglets are used
to simulate the response of infants, then perhaps the theory
that foals can also be compared to piglets has some
validity.
The use of the pig as a model has both advantages and
limitations. Being able to do research on piglets that are
littermates and on pigs from different litters, but by the
same parents, allows experiments to be performed using large
numbers of animals with minimal genetic variation. Also,
many animals can be studied in a relatively short period of
time, as the gestation of the sow (approximately 114 d) is
much less than that of the mare (approximately 335 d). Sows
can be bred to farrow all year, while mares have a foaling
season of several months.


7
Digestive Problems of Neonates
After birth, mammalian neonates undergo a period of
rapid acclimation to extrauterine life. They must adapt to
using enterally rather than parenterally supplied nutrients.
The neonate is now a free-living organism and it must be
able to ingest and absorb nutrients from its environment.
In mammals, of course, this usually means learning to nurse
from its mother. Serious problems occur when the newborn
has difficulty suckling or has an incompetent digestive
tract. Immature or damaged intestinal tissues will not
function normally, therefore, it is necessary to consider
the nutritional needs of neonates with digestive problems
separately from those of healthy neonates.
Diarrhea is a potentially serious problem for neonates
and can occur for several reasons. The causes are usually
classified as infectious or non-infectious.
Infectious Causes of Diarrhea
Diarrhea in piglets and foals has been attributed to an
assortment of viruses and bacteria. Rotavirus, in
particular, seems to be present in over 30% of the cases,
while coronavirus and adenovirus are also common infectious
agents (Palmer, 1985; Bergeland and Henry, 1982). These
authors suggest that rotavirus acts by causing damage to the
epithelium of the small intestine and leaving the neonate
vulnerable to a secondary bacterial infection. The bacteria


8
usually cultured in cases of piglet diarrhea include
Salmonella, and E. coli, while the latter is not considered
a problem in foals. They also suggested that in foals,
bacterial diarrhea may be secondary to septacemia and
enteritis. Piglets may also be plagued by coccidiosis or
infestations of strongyles.
Because of the ever present threat of invasion by
infectious organisms, the area where newborns are housed
must be kept extremely clean. It has been suggested that
foals usually become infected through contact with adult,
asymptomatic shedders on the same farm. Treatment for
diarrhea in piglets and foals usually includes
administration of antibiotics to cause a change in gut
flora, supplemental vitamins to offset decreased gut
absorption and bacterial synthesis, and, in piglets,
antispasmodics (Mouwen, 1971; Palmer, 1985).
Non-Infectious Causes of Diarrhea
Stress of all kinds, including exposure to low ambient
temperatures, or malfunctioning of the small intestine can
predispose newborn animals to diarrhea (Bergeland and Henry,
1982) .
Many foals suffer from diarrhea on d 6 to 14
postpartum. Because this is also the time when the mare
returns to estrus after giving birth, it has been called
"foal heat" diarrhea. However, there was no difference in


9
the amount or severity of diarrhea in hand-fed or mare-fed
foals observed by Palmer (1985). Therefore, the theory that
hormonal changes in the mare are reflected in the milk is
unfounded. It is important, though, to monitor the foal
closely at this time, because the onset of infectious
diarrhea can mimic foal heat diarrhea.
Piglets may excrete thin, watery feces called "white
scours" when they are very young. They usually seem normal
in attitude, but may be less active. The feces, upon
analysis, usually contains greater than 50% fat, but it is
not known if this is the cause or effect of the condition.
One experimenter suggests that it is the presence of
abnormally structured gut villi that is responsible for the
diarrhea (Mouwen, 1971).
Lipid Metabolism in Neonates
Fat in the Neonatal Diet
The addition of fat to human and animal enteral
formulas has proven to be an effective means of increasing
caloric density in an easily digestible form without also
increasing osmolarity (Koterba and Drummond, 1985). By
increasing the caloric density, the volume of formula to be
fed can be kept low. Large fluid loads can be detrimental
to sick or premature newborn patients. Fats are also added
to enteral and parenteral solutions to provide essential


10
fatty acids and to provide a carrier for fat soluble
vitamins.
Interest in studying the type of fat most readily
utilized by neonates has increased because the composition
of fat in fetal pig tissues and in sow's milk and colostrum
can be influenced by the source of dietary lipids fed to the
sow. Therefore, it may be possible to manipulate the diet
of piglets to affect the compostion of body lipids. This is
important because there is evidence that piglets metabolize
fats differently according to chain length and degree of
saturation (Miller et al., 1971; Lloyd and Crampton, 1957).
This also appears to be true in human infants (Faber et al.,
1988; Bach and Babayan, 1982; Lammi-Keefe and Jensen, 1984).
The fat content of milk and the extent of body lipid
stores at birth vary with species. There is usually an
increase in body fat of young animals after nursing.
Neonates who are energy-deprived must depend on fat reserves
for metabolizable energy and for thermal regulation
(Pegorier et al., 1983).
It has been suggested that within 2 d of birth neonatal
piglets can utilize fat as efficiently as glucose and the
ability to use fat increases with age (Miller et al., 1971;
Wolfe et al., 1978). Piglets have low fat stores at birth
(approximately 1% of their body weight) that rapidly
increase to about 15% of their body weight by day 14.
Wolfe et al. (1977) fed isocaloric diets to colostrum-


11
deprived, newborn piglets. The diets differed only in the
proportion of calories provided by carbohydrate and
butterfat. The fatty acid pattern of the backfat of the
piglets resembled the fatty acid composition of the diet,
especially when high fat diets were fed. Lipogenic enzyme
activity decreased with higher fat diets, which the authors
suggest is due to the inhibiting effect of increased lipid
intake rather than to decreased carbohydrate intake. Wolfe
and coworkers in 1978 determined that on low fat diets,
newborn piglets will resort to de novo fat synthesis and
attain a level of fat deposition similar to piglets fed a
high fat diet.
Fatty Acid Digestion and Absorption
Researchers have investigated the digestion and
absorption of lipids in neonates of many species, but more
information on how different types of fats are utilized by
premature and compromised newborns is needed. Data on lipid
metabolism in foals are limited.
Of particular interest is the effect of chain length on
neonatal absorptive and metabolic processes. In the past,
infant formulas contained exclusively long chain
triglycerides (LCTs), but recent work suggests that
compromised neonates may benefit from the replacement of
some of the LCT with medium chain length fats. It has been
suggested that medium chain triglycerides (MCT), usually


12
saturated fats consisting of 6 to 12 carbons, are readily
hydrolyzed and absorbed by infants, rat pups, and piglets
(Newport et al.; 1979, Putet et al., 1987; Jandacek et al.,
1987).
Medium chain fatty acids (MCFA) from the diet are
transported directly to the liver via the portal vein
whereas long chain fatty acids (LCFA) travel through the
lymph and then through the general circulation before
reaching the liver (Newport et al., 1979). MCT being more
polar and therefore more water soluble than LCT, are more
easily attacked by pancreatic lipase and thus, are a
desirable alternative to LCT in cases of pancreatic
insufficiency or in other situations when LCT would be
poorly absorbed (Jandacek et al., 1987).
Specifically designed triglycerides have been used to
benefit from both MCT and LCT in a single lipid source. A
triglyceride having medium chain fatty acids at the one and
three positions and a long chain fatty acid at the two
position may be the solution. This takes advantage of the
pancreatic lipase specificity for cleavage at the one and
three sites, leaving a 2-monoglyceride. Even in cases where
LCT absorption is poor, the 2-monoglycerides seem to be
easily transported across the intestinal mucosa (Newport et
al., 1979), thereby avoiding essential fatty acid (EFA)
deficiency. Human milk triglycerides have a 16:0 at the 2


13
position, which is thought to enhance the absorption of
dietary fats (Lammi-Keefe and Jensen, 1984).
Metabolism of Medium Chain and Lona Chain Fats
bv Neonates
The different metabolic pathways available to MCFA and
LCFA affect the energy sources available to peripheral
tissues. Interest in substituting MCT for LCT as part or
all of the lipid content of enteral formulas stems from the
products of metabolism of MCT. Several groups of
investigators have begun feeding MCT to neonates in an
effort to determine its benefits and drawbacks.
There is evidence to suggest that infants, especially
preterm infants, do better on MCT-added formulas than on
cow's milk alone, even though MCT are not found in large
amounts in mature human milk (Putet et al., 1987). Lammi-
Keefe and Jensen (1984) found considerably more 10, 12, and
14 carbon fatty acids in the milk of mothers that had very
premature or premature babies when compared to the milk of
mothers with full-term babies. They speculate that this may
be of some special benefit to premature infants. Some
researchers advocate feeding MCT to infants that have
malabsorption of lipids (Bach and Babayan, 1982).
MCT and MCFA are not normally found in the blood even
when they are added to the diet. They are thought to be
rapidly metabolized by the liver and do not reach the


14
peripheral tissues intact. However, 15-20% of the fatty
acids found in the umbilical cord blood of infants have
eight carbons or less. MCT tend to increase carbohydrate
tolerance, insulin secretion, and decrease glucose output
from the liver when given orally (Bach and Babayan, 1982).
A study on neonatal puppies (Cotter et al., 1987)
suggests that insulin causes LCFA to be removed from the
blood for storage, while MCFA are removed for immediate use
as an energy substrate. The authors have shown that low
doses of MCT given intravenously (i.v.) are utilized more
rapidly than LCT. The calories are available faster because
the lipoprotein lipase (LPL) in the capillary walls quickly
releases the MCFA, which then bind to albumin and are
absorbed by peripheral tissues. In the liver, MCFA can
enter the mitochondria rapidly, in contrast to LCFA which
must be activated and be transported with the help of a
carnitine carrier. MCFA can be oxidized so quickly that the
Kreb's cycle is overwhelmed. When this happens, the 2-carbon
units are funneled into ketone bodies which then diffuse
into the blood. The authors suggest that MCT is a more
flexible energy source than LCT. They also observed an
increase in plasma clearance rate of long chain fats when
medium chain fatty acids were present.
In a similar study, piglets fed either a 25:75 or 50:50
mixture of dietary MCT and LCT had higher blood glucose
concentrations and increased insulin secretion than pigs fed


15
a diet in which the fat source was 100% LCT (Wolfe et al.,
1978) .
Newport et al. (1979) fed 2 day old neonatal pigs
various ratios of MCT and LCT until they were euthanized at
day 28. The pigs fed a 50:50 ration had similar growth
rates to pigs fed 100% LCT as a dietary fat source.
However, 90% MCT feeding resulted in a significantly slower
growth rate. The pigs fed 90% or 50% MCT had higher liver
weights but lower carcass weights than the 100% LCT group.
The addition of MCT resulted in lower plasma lipid and free
fatty acids (FFA), but higher body water, total body lipids,
and cholesterol. They found no increase in nitrogen
retention due to MCT feeding as had been suggested by
others.
MCT feeding is not without possible disadvantages.
Urinary output of dicarboxylic acids (DCA) may be seen.
These end products of omega oxidation of medium chain fatty
acids, such as seberic, sebacic, and adipic, are often
detected in the urine during fasting, during ketosis, or in
children with defects in fatty acid oxidation or carnitine
metabolism. Henderson and Dear (1986) fed preterm infants
averaging 29 wk gestational age either parenterally (no
medium chain fats included) or enterally. The enteral
formulas contained either a high amount of MCT (1.8g/100ml)
or a low amount (.44 g/100 ml). Some infants received
breast milk containing .15 g MCT/100 ml. The infants fed


16
the high MCT formula excreted larger amounts of DCA than
infants fed the other feeding regimens, although no other
symptoms were evident. It is unclear if this would be a
problem for healthy infants. The authors stated that MCT
feeding would be unadvisable in infants with 0-oxidation
defects and that more investigation is needed to determine
the effects, if any, of increased urinary excretion of DCA.
They did not comment on the possibility that DCA excretion
resulted from inefficient use of MCT by the infants.
Ketones from MCT provide a guick source of energy for
neonatal extrahepatic tissues. As their concentration in
the blood increases, utilization by extrahepatic tissues
increases. Therefore, they are a good substrate for energy
production in times of high energy needs, such as growth or
undernutrition (Bach and Babayan, 1982). Ketogenesis is
controlled by a feedback system: when the concentration of
ketones in the blood increases, negative feedback to the
liver decreases production (Robinson and Williamson, 1980).
Gentz et al. (1970) have shown moderate increases in
plasma ketones during starvation in newborn and 16 day old
pigs during times when fat utilization is limited. They
suggest that this may not be detrimental because 10 kg pigs
are not prone to ketosis during starvation. Pegorier et al.
(1983) question the extent of ketogenesis and 0-oxidation in
newborn pigs since they do not show hyperketonemia as do
rabbits, rats, guinea pigs, and humans fed high fat diets.


17
In summary, the MCT system has a fast turnover rate but
is easily overloaded. The presence of medium chain fats
stimulate the absorption of plasma FFA. The 2-carbon units
resulting from /3-oxidation of MCFA are ketogenic, thereby
providing the flexibility of an alternative energy source.
The LCT system handles large quantities well but has a slow
turnover rate. The resulting increase in plasma FFA may
lead to acidosis. Some LCT are needed in the diet to
prevent essential fatty acid deficiency. MCFA may improve
the utilization and absorption of LCFA when there is
impaired lipid digestion.
Disaccharidase Activity in Neonatal Small Intestine
Introduction
Milk is a complex mixture of nutrients, hormones, and
other substances designed to provide optimal nutrition for a
normal, healthy newborn. Consequently, it is a difficult
task to formulate an artificial diet that will allow the
hand-raised neonate to grow and develop normally. The
challenge is even greater when the neonate is sick, weak or
premature.
The developmental pattern of intestinal disaccharidases
has been used to assess the nutritional status of neonates.
It has been shown that improper nutrition results in a
malnourished intestinal tract that does not show the same
pattern of enzyme activity changes as a healthy neonate


18
nursing its mother (Rossi et al., 1986). These authors
experimentally malnourished rats adding extra pups to a
normal-sized litter. The pattern of disaccharidase activity
in the malnourished rat pups resembled that of lesser
developed pups. The maltase and sucrase activities were
lower, and the lactase activity higher than in the control
pups.
Sick neonates often lose the ability to digest lactose
(Tzipori et al., 1984) and in infants, prematurity may
result in low total intestinal lactase activity (Mayne et
al., 1986). Therefore, it is important to assess the value
of alternative carbohydrates when developing artificial
formulas Dairy farmers are also interested in the use of
economical energy sources for use in calf milk replacers
(Dollar and Porter, 1957). The efficient utilization of any
energy source will depend on the activity of appropriate
intestinal enzymes for digestion and absorption. Digestion
of complex carbohydrates, such as corn starch, are cleaved
by amylases into smaller units, into maltose, and finally
into single glucose molecules that can be absorbed (Code,
1968). Therefore, starch can not be utilized unless there
is adequate amylase and maltase present. Roberts (1974)
detected only low levels of both heat-sensitive pancreatic
a-amylase and heat-stable glucoamylase in neonatal foals.
One way to determine the digestibility of various sugars for


19
use in neonatal formulas is to estimate the amount of
disaccharidase activity present in the intestine.
Brush Border Disaccharidases
Disaccharidases are synthesized in membrane-bound
polysomes within enterocytes of the small intestine. There
is evidence to show that they are present in an inactive
form in crypt cells, becoming activated when the cells have
migrated to the top of the villus. Active lactase enzyme
has only been found in the microvillus membrane on "mature"
enterocytes at the tip of the villi in neonatal rats
(Mackenzie, 1985). The enzymes are activated when proteases
cleave a polypeptide chain from the large pro-disaccharidase
molecule. Extracellular pancreatic proteases are
responsible for activating maltase and the
sucrase-isomaltase complex, while lactase is activated by
intracellular proteases. The disaccharidases are attached
to the brush border of the epithelial cells lining the
intestine. In the rat, the rate of turnover for these large
proteins is short, about 11.5 h. The enterocytes themselves
take only 4 to 5 d to migrate from the crypt to the tip of
the villus, where they are shed (Dahlqvist and Semenza,
1985). The rapid rate of cell division and protein
synthesis makes these cells and their attached enzymes very
susceptible to adverse conditions and poor nutrition.


20
Disaccharidase enzymes, primarily sucrase, maltase, and
lactase, tend to follow the same pattern of development in
most mammals, but the rate of development, pre- and post
partum, differs by species (Paige and Bayless, 1981). The
amount of each disaccharidase present in the intestine will
determine the carbohydrates that can be successfully
incorporated into milk replacer formulas. Milk contains
many substances that can affect the development of the
intestines. Therefore, artificial formulas must be able to
support the normal pattern of maturation seen in neonates
nursing their mother. Determination of enzyme activity will
allow assessment of the development and general health of
the intestine. In this way, the nutritional value of
artificial diets can be compared to that of milk and
colostrum.
Developmental Patterns of Enzyme Activity in Neonates
The relative progression of the increase and decrease
in disaccharidase enzymes is similar for many animals, but
the timetables are very different for each species. For
most mammals, lactase is high at birth, increasing during
the nursing period, and decreasing near weaning. Lactase,
which develops in human infants later in gestation than
sucrase and maltase, reaches a peak at 36 to 40 wk gestation
(Roggero et al., 1986). Therefore, very premature infants
will have a limited ability to digest this sugar. Birth and


21
feeding each contribute to a postnatal rise in lactase in
both term and preterm infants (Weaver et al., 1986).
Studies on neonates of several species indicate
that prematurity or damage to the intestines can greatly
reduce lactase activity. Mayne and coworkers (1986)
developed a technique for assessing disaccharidase activity
in living human infants. They demontrated a significant
correlation between the enzyme activities measured in
jejunal fluid with enzyme activities in adjacent mucosal
tissue. Using this method, they studied eight premature,
but appropriate for gestational age infants 17 times during
the first 3 wk after birth. The infants were fed
pasteurized breast milk via nasogastric tube. The lactase
activity in these infants at 1 wk of age was lower than the
activity present at 2 and 3 wk of age (P < .001). The 1 wk
old infants had enzyme activities in the normal range for
adults. This work agreed with other published reports
indicating that lactase activity increases greatly near the
end of gestation in humans. This results in a lowered
capacity of preterm infants to digest a lactose-
based diet.
Damage to the intestinal tissue may also result in
reduced lactase activity. Biopsies were obtained proximally
and distally to a complete or incomplete congenital blockage
of the small intestines of 12 newborn infants during surgery
(Serrano and Zetterstrom, 1987). The activities of all


22
disaccharidases in tissue samples proximal to the
obstruction site were reduced when compared with previously
published reference values. Lactase activity was more
markedly reduced than the activities of the other
disaccharidases. There was no difference in lactase
activities in tissues from the four infants with complete
blockages compared the eight infants with incomplete
blockages.
However, when tissue samples from sites distal to the
obstruction were compared, lactase activity was lower (P <
.03) in infants with complete blockage when compared with
infants with incomplete blockage. The authors suggest that
distension and damage to the small intestine proximal to the
blockages resulted in impaired development of brush border
enzymes, particularly lactase. Minimal passage of ingesta
through an incomplete blockage permitted greater development
of lactase activity than detected in tissue distal to a
complete blockage.
To investigate the effect of infectious pathogens on
disaccharidase activities in neonates, Tzipori et al. (1984)
inoculated seven 4 day old foals with various combinations
of Streptococcus durans, Staphylococcus epidermis, and
Escherichia coli isolated from feces of foals with naturally
occurring diarrhea. Three foals received saline
inoculations. Inoculations were adminstered via
nasogastric tube and were followed with 200 ml evaporated


23
cow's milk. The foals were necropsied 1 to 3 d later. The
seven foals inculated with pathogens had experienced
diarrhea. S. durans bacteria were found to be adhered to
the intestinal mucosa. This probably contributed to the
significantly lowered lactase activity measured in these
foals compared with the lactase activity of the control
foals. These authors then conducted a similar study on
newborn piglets, inoculating them with the same bacterial
strains isolated from foals. These piglets also experienced
diarrhea and reduced lactase activity compared with saline
inoculated controls.
Maltase and sucrase do not appear in any appreciable
amount until at least a few days after birth (Code, 1968;
Veum and Mateo, 1986; Manners and Stevens, 1972; Ahrene et
al., 1969). This is the case for rabbits, calves, pigs,
rats, dogs, and cats (Code, 1968; Paige and Bayless, 1981).
Guinea pigs, however, are very mature at birth, nurse
infrequently, and can digest maltose and sucrose immediately
(James et al., 1987). Human infants develop considerable
maltase and sucrase activity before birth (Paige and
Bayless, 1981).
It has been reported that newborn piglets can utilize
lactose and glucose equally well at birth (Dollar et al.,
1957; Ahrene et al., 1969), while sucrase and maltase
activities increase during the first few days of life (Veum
et al., 1986). Dahlqvist (1961a) reported high lactase


24
activity in the pig at birth and a rapid increase in maltase
by day 2, but he found no appreciable sucrase at this age.
In contrast, Beilis (1957) reported that by day 3, piglets
could digest sucrose, lactose, and glucose equally well.
James et al. (1987) found low levels of maltase and sucrase
in sow-fed piglets on d 1 to 4 of life, with levels
increasing from day 3 through day 10. Sucrase activity was
twice that of maltase. When they gave Epidermal Growth
Factor (EGF), which may be present in colostrum, to
hand-raised piglets, they found an increase in sucrase and
maltase in the middle and distal portions of the small
intestine. EGF had no effect on lactase.
Dairy calf producers have tried to formulate milk
replacers using economical energy sources other than
lactose. Dollar and Porter (1957) fed newborn calves skim
milk using starch as an energy source with and without
amylase. Calves under 3 wk of age did poorly on this diet.
The authors suggest that the calves were unable to digest
the dextrins and maltose resulting from amylase digestion.
They report only very low activities of maltase and amylase
in the newborn calf and no sucrase activity. They state
that calves under 4 wk old have only been shown to digest
lactose and glucose. They did not see an appreciable rise
in maltose digestion until the calves were 9 wk old.


25
Distribution of Disaccharidases
The distribution of disaccharidase activity in the
small intestine has been investigated in several species.
In calves, lactase activity is highest in the proximal small
intestine. With age, lactase activity is reduce, mostly in
the distal third (Huber et al., 1961). Manners and Stevens
(1972) studied enzyme distribution in the newborn pig.
While they found great variation from pig to pig, they were
able to draw some conclusions. Enzyme activities were
measured at various sites along the small intestine, with
the proximal and distal ends represented as the 0% and 100%
sites, respectively. Lactase activity was highest at the
20% site along the small intestine, decreasing to a low at
the 95% site. From birth to 1 wk old, much of the decrease
seen in lactase activity occurred in the proximal 50%, with
higher activities found in the more distal section. Sucrase
activity was only barely detectable until the pigs were 1 wk
old, with the highest values in the proximal two-thirds of
the small intestine.
In foals, lactase activity was reported to be highest
in the duodenum and upper jejunum, decreasing distally
(Roberts et al., 1974). Maltase activity was low in the
equine fetus, increasing slowly during the first month after
birth mostly in the proximal and middle segments of the
small intestine, then increasing markedly until the end of
the first year. Sucrase was very low at all sites along the


26
intestine in the newborn foal, increasing in the proximal
sections during the first year, with the pattern of
distribution parallel to that of maltase.
Postmortem studies of the entire length of small
intestine of preterm and near-term human infants provided
information on the development of disaccharidases (Raul,
1986). The relative distribution of sucrase and lactase was
similar for infants in both age groups throughout the
intestine. The greatest amount of lactase activity was
found in the jejunum and duodenum, with little found in the
distal small intestine. Term infants had five times greater
lactase activity than the preterms, with the largest
differences found in the proximal intestine. The greatest
amount of sucrase activity was found in the jejunum. The
term infants had more enzyme activity in all segments of the
intestine. Glucoamylase was present in the tissues of
preterm infants, increasing distally, as it does in adults.
The authors suggest that this enzyme, along with salivary
amylase (which, in preterm infants, is still functional in
the small intestine) enhances digestion of glucose polymers.
Sucrase
Veum and Mateo (1986) fed pigs a sucrose-added formula
but could not stimulate sucrase activity by day 7. No
appreciable amounts of the enzyme were detected until day
14. Manners and Stevens (1972) found that pigs fed an


27
artificial diet had higher sucrase activity than sow-fed
controls. This may suggest that diet influences the rate of
development of the intestine. Huber et al. (1961) reported
that even when calves were fed a sucrose-added diet, there
was no notable digestion of sucrose until day 44, in
contrast to their study with pigs, where sucrase digestion
was evident by day 10.
Very low sucrase activity was detected in the small
intestine of equine fetuses and newborn foals. The activity
increased slowly during the first months after birth,
reaching adult activities by 7 mo of age (Roberts, 1974).
Lactase
Humans are an unusual species. For some races, the
ability to digest lactose remains long after weaning, but
the percent of the population with adult lactase activity is
small (Dahlqvist and Semenza, 1985). In humans, rabbits,
and rats prolonged feeding or absence of feeding of lactose
does not affect the postnatal changes in lactase activity
(Paige and Bayless, 1981). In calves, up to 44 d of age, no
increase in lactase activity occurs as a result of adding
lactose to the diet (Huber et al., 1961).
In human infants, lactose hydrolysis by the
/3-galactosidase enzyme is affected by other mono- and
disaccharides that may be concurrently ingested (Paige and
Bayless, 1981).


28
Lactase activity in the neonatal piglet has been shown
to be high at birth, reaching a maximum at day 15, then
decreasing throughout the nursing period with low activities
still detectable in the adult (Paige and Bayless, 1981; Veum
and Mateo, 1986; Manners and Stevens, 1971; Dollar et al.,
1957; Dahlqvist, 1961b).
Lactase activity was detected in the mucosa of eguine
fetuses as early as 105 d of gestation, increasing after the
ninth month, reaching a maximum at birth and remaining high
for 4 mo (Roberts et al., 1974). The activity declined,
especially in the duodenum, until 1 yr of age.
Maltase
Several researchers have investigated maltose as an
energy source for neonates. Cunningham and Brisson (1957)
fed 1 day old piglets a purified casein-based diet with
either maltose or glucose as an energy source. In piglets 3
to 7 d old, greater than 95% of the maltose in the diet, and
greater than 99% of the glucose disappeared from the small
intestine. The glucose diet resulted in greater
digestibility for protein and carbohydrate than the maltose
diet (P < .01). When the entire small intestine of neonatal
piglets of various ages was ligated and injected with a 5%
maltose solution, .66 /mol of maltose was hydrolyzed and
absorbed per h per kg body weight. This increased to 1.05


29
jLiinol by day 6 and 7. The authors conclude that maltose
digestion during the first wk may only be borderline in
supporting the animal's energy needs, but after day 7
maltose is used just as efficiently as glucose. Huber et
al. (1961) could not increase the maltase activity of calves
by feeding diets of whole milk plus lactose, or sucrose and
starch for 4 4 d. They note that this is in contrast to
other workers who have reported an increase in maltase
activity beginning in the third or fourth wk after birth.
Using intestines obtained from slaughterhouses, Roberts
et al. (1974) detected low maltase activity in equine
fetuses during the first 3 mo of gestation. Activities
remained low until birth. Enzyme activity doubled during
the first month, with the greatest increase occurring in the
proximal small intestine. Adult activities were achieved by
7 mo of age. No data were given specifically for the early
neonatal period.


30
Determining Disaccharidase Activity in Experimental Animals
Intestinal disaccharidase activity can be determined by
homogenizing samples of tissue from segments along the small
intestine and incubating the homogenate with disaccharide
substrates. The glucose produced is a measure of enzyme
activity. Many researchers have used this method, as
modified from Dahlgvist (1964). The major disadvantage of
this method is that the experimental animal must be
euthanized. Economic constraints usually limit the number
of foals that can be euthanized, making it impossible to
conduct large scale equine trials of this type. Therefore,
it would be helpful to have an alternate method, at least
during preliminary trials. Studies on calves have shown
oral tolerance tests to correlate well with the results
obtained using tissue homogenates when comparing relative
enzyme activities (Dollar and Porter, 1957).
Roberts (1975) used an oral disaccharide tolerance test
to determine the digestibility of various sugars in adult
horses and ponies. Although foals were not included in the
study, Roberts suggested that an oral lactose tolerance test
would be a useful tool for determining digestive competence
in the sick or premature foal. Testing the digestibility of
other sugars (for possible inclusion in artificial formulas)
in this way would allow animals to be studied without
euthanasia.


31
Responses of the Neonate to Stress
Stress factors include a wide variety of situations
that can cause problems for neonates. Poor maternal
nutrition and dystocia are examples of stresses can occur
before or during birth, but this project will be concerned
with stresses that occur during the neonatal period.
Failure to Acquire Passive Immunity
In the horse and pig there is little transport of
immunoglobulins across the placenta. So, these species must
depend on colostral transfer of antibodies to protect them
from a wide variety of infectious organisms until their own
immune systems are competent. The risk is high that these
neonates will be severely stressed by infections if they are
deprived of colostrum or are unable to absorb antibodies
during the first day of life. This becomes even more
serious if the neonate is also exposed to other stresses in
its environment or is weak, sick, or immature at birth.
Many researchers have emphasized the importance of
colostrum ingestion for newborn foals and have suggested
reasons why foals may be hypogammaglobulinemic even after
they have nursed (Naylor and Bell, 1985; Naylor, 1979;
Rossdale, 1985). Their findings show that immunoglobulins
in the colostrum decline rapidly, becoming 15% of its
concentration at parturition in 4 to 8 h.


32
Mares may produce poor quality colostrum for several
reasons. Older broodmares may leak colostrum before
parturition and so lose the high antibody content necessary
to provide the foal with adequate protection. Some foals
are born prematurely, before the mammary gland has completed
concentrating the immunoglobulins. Foals must ingest
colostrum within 24 h of birth. After this time, the
digestive tract "closes" and will no longer absorb large
molecules. Closure is not delayed by food-withholding.
Foals may be deprived of colostrum if they are orphaned
or because of a problem with the mare or with the foal
itself that prevents it from nursing. Foals that need to be
hospitalized may be separated from their mothers. Frozen
colostrum may either be unavailable or unwise to feed,
depending on the condition of the foal. Colostrum-deprived
foals have been shown to remain almost completely
agammaglobulinemic for 2 wk, then begin rapid synthesis of
antibodies until they appear the same as colostrum-fed foals
by 5 wk of age (Naylor, 1979). These workers feel that over
50% of colostrum-deprived foals will develop an infection
resulting in septacemia, diarrhea, pneumonia,
omphalophlebitis, or arthritis within 1 wk of birth. They
suggest prophylactic treatment of 1.2 liters of plasma taken
from an adult horse, given intravenously over 2 h, to supply
an adequate amount of IgG.


33
Piglets are also likely to suffer similar diseases if
not provided with passive immunity from colostrum (Svendsen
and Bille, 1981; Bergeland and Henry, 1982) or kept in
isolated, aseptic conditions (Pond, 1978). In addition to
providing passive immunity to neonates, colostrum also
contains epidermal growth factors that promote mucosal
development in the gastrointestinal tract (Svendsen and
Bille, 1981). Unfortunately, it has been shown that
colostrum-deprived neonates fed either milk from other
species or soy-based formulas sustain damage to the
intestine, as the immune system responds to the foreign
proteins ingested (Weaver, 1986). The use of elemental
diets would prevent this adverse reaction as they contain
hydrolyzed rather than whole proteins.
Cold Stress
Exposure to a low ambient temperature shortly after
birth is a common problem for many neonates including
piglets, foals, and human infants. Management practices
used on many breeding farms encourage mares to begin cycling
and ovulating earlier in the year than they would in nature,
so foals are born more and more frequently in late
winter/early spring when temperatures are low. Therefore,
it is important to study the ability of neonates to regulate
their body temperature and the effects of cold stress on
their health and nutritional requirements.


34
Normal body temperature for foals is 38.05C 1, with
the lower boundary of the thermal neutral zone being 14C
during the first 48 h postpartum. After that, the foal can
tolerate an ambient temperature as low as 8C, provided he
has nursed and the environment is free of draught (Rossdale,
1985). Premature foals often have difficultly in
maintaining their body temperature, which may drop to 36.6C
or even lower (Naylor, 1979). Foals lack significant
deposits of brown adipose tissue and must rely on shivering
for thermogenesis (Rossdale, 1985).
The normal body temperature of piglets is 39C at
birth. The thermal neutral zone is very small (34-35C)
(Stanton et al., 1973) and they have greater difficulty
maintaining their body temperature under cold stress than do
foals and calves (McCance and Widdowson, 1959) .
Piglets are easily cold stressed for several reasons:
they are poorly insulated at birth, having almost no brown
adipose tissue and only 1% of their body weight (about 10 g)
is fat; they have a small body size with a large surface
area; their metabolic rate is low at birth; and they are
born with no protective hair or fur (McCance and Widdowson,
1959; Gentz et al., 1970; Curtis and Rogler, 1970; Stanton
et al., 1973). However, their ability to thermal regulate
does increase with age and is fairly well developed by 1 wk
of age (Gentz et al., 1970; Curtis and Rogler, 1970; Stanton
et al., 1973) .


35
Piglets will experience a 2 degree decrease in body
temperature, to 37C, shortly after birth. Adverse effects
may be seen if the temperature reaches 35C (Svendsen and
Bille, 1981).
An ambient temperature of 5C is close to the cold
limit for newborn pigs. At this temperature, there should
be maximum utilization of their thermal regulatory
capabilities. Several investigators have exposed newborn
pigs to this temperature for short periods of time. Results
suggest that starving piglets undergoing cold stress
experience hypoglycemia and will not survive longer than 30
h unless fed (McCance and Widdowson, 1959). These workers
report that at birth, pigs are able to increase oxygen
consumption during cold stress whether they are fed or not,
whereas the newborn rat will not show an increase until it
is 5 d old. Rabbit young improve their ability to respond
during the first few days of life.
Carbohydrate is the main energy source of newborn
piglets, even though sow's milk contains 30-40% fat on a dry
matter basis (Allee et al., 1971) because fat utilization is
limited during the first few days postpartum, regardless of
the ambient temperature. However, cold exposure results in
rapid depletion of energy stores and death from hypothermia
unless the piglets are fed. It is estimated that the
carbohydrate and fat stores endogenous to the piglet would


36
yield about 72 h worth of energy with ambient temperatures
in the thermal neutral zone (Gentz et al., 1970).
Compromised Neonates
Being born premature or dysmature will compromise the
neonate's ability to adapt to extrauterine life. Dystocia
and neonatal maladjustment syndrome (where there is reduced
ability to suck, swallow, or move about) also predispose
newborns to septacemia and in the case of foals, septic
arthritis (Koterba et al., 1984). Premature foals usually
have abnormal blood gases and an impaired acid/base status
(Rossdale et al., 1987). They often are weak and take a
longer-than-normal time to stand and nurse. They commmonly
show hypoflexion of the metacarpal and metatarsal joints,
and a bright red tongue, but they may maintain normal
respiration and heart rate for the first 24 h before their
condition deteriorates (Rossdale, 1987).
Intestinal maturation will not be complete in preterm
neonates. Enterocyte function, villi development, and
enzyme activity are often different from that found in term
newborns, resulting in decreased digestibility of some
nutrients (Raul et al., 1986; Mayne et al., 1986; Mackenzie,
1985).


37
Hematology and Blood Chemistry of Neonates
Hematology
It has been suggested that evaluating the packed cell
volume (PCV), hemoglobin (Hb), red blood cell count (RBC),
white blood cell count (WBC), and differential leukocyte
count can be helpful in assessing the condition of neonates
and estimating the degree of maturity and viability of
neonates (Rossdale, 1985).
Becht and Semrad (1985) found that in foals, PCV, Hb,
and RBC counts peak at birth and then begin to decrease
within 12 h postpartum. They also noticed that mean
corpuscular volume (MCV) decreased slightly in the fetus
just before birth, then remained steady for the first 2 wk
of extrauterine life. They suggest that some of these
indices may be useful for establishing the maturity of the
foal at birth.
Several workers have reported lower leukocyte counts (4
x 10 /liter) for premature foals as compared to full term
Q
foals (6 x 10 /liter). Erythrocyte counts were also lower (6
x 1012 vs 12 x 1012/liter) as was the PCV (33 vs 45.6%).
Globulin (<10 gm/liter) and gammaglobulin (4 gm/liter) were
also lower than for normal foals (Jeffcott et al., 1982;
Kitchen and Rossdale, 1975; Rossdale, 1983; Becht and
Semrad, 1985).
Foals have an intact granulocytic system at birth. The
increase in WBC reported during the first day of life is


38
thought to be due to an increase in mature polymorphonuclear
neutrophils (PMN) and lymphocytes. The increase in PMN
continues during the first 3 days postpartum. During the
first wk, monocytes begin to appear. Basophils are not
normally seen in the neonatal period.
9
Premature foals tend to have lower WBC (4 x 10 /liter) ,
RBC (6 x 1012/liter), PCV (33%), globulin (<10gm/liter),
gammaglobulin (4gm/liter) and a narrower neutrophil to lymph
ratio (1.2:1) than normal foals. A severe leukopenia, left
shift, and the appearance of toxic neutrophils may indicate
sepsis (Becht and Semrad, 1985).
Jeffcott et al. (1982) and Rossdale (1983) found that
premature foals consistently show a lower than normal
neutrophil to lymphocyte ratio (N/L). Normal foals have a
ratio > 2:1 while premature foals show a reversed N/L ratio
of > 1:1. A dose of short acting exogenous ACTH will elict
a neutrophilic response in mature foals but not in immature
foals that lack normal adrenal function.
In piglets, RBC (from 6.18 x 1012/liter to 4.4 x
1012/liter), PVC (from 40 to30%) and Hb (12.5 to 10.0g/dl)
are decrease during the first wk after birth. However, the
number of WBC tends to increase from about 6.2 to 17 x
3
10 /n1. The proportion of PMN to lymphocytes percentages
shifts from about 38:60 to about 53:42. Granulocytes are
rarely seen during the first wk (Schmidt and Tumbleson,
1985) .


39
Chemistry
Various clinical chemistry analyses are useful for
evaluating the general health and nutritional status of
newborn animals. Both foals and piglets are susceptible to
hypoglycemia shortly after birth, especially if they are
weak, sick or stressed.
Normal blood glucose in the neonatal piglet is 60 to 80
mg/dl. However, it may be as high as 100 mg/dl immediately
after birth. The decrease may be due to limited glycogen
stores in the liver (Pond, 1978). Blood glucose values as
low as 48 mg/dl have been reported for newborn pigs,
increasing to 114 mg/dl by day 7.
Foals are susceptible to hypoglycemia shortly after
birth, especially if they are weak, sick or stressed. Foals
that have suckled during the first 2 h after birth were
reported to have the following blood glucose values: 95.5 +
17.44 mg/100 ml at birth, 83 3.28 mg/100 ml 30 min
postpartum, and 131 + 12.4 by 14 h (Kitchen and Rossdale,
1975). Normal, healthy foals that have suckled have blood
glucose levels that are higher than adult levels for the
first 24 h and remain in the high normal adult range for the
first 30 d. Hypoglycemia may result in sick foals not
receiving sufficient nutrient intake, or from poor digestive
and absorptive function even with adequate intake (Becht and
Semrad, 1985).


40
Blood lipid components such as free fatty acids,
trigycerides, ketones, blood urea nitrogen (BUN), and
circulating liver enzymes are important parameters to
consider when evaluating the utilization of dietary fat and
nitrogen status in the neonate.
Milk Composition and Intake
Some researchers, using milk replacers currently
available for foals, have expressed dissatisfaction with the
foals' growth rate and general appearance. They report that
foals fed milk replacers according to directions usually
remain small (Naylor and Bell, 1985). Since no other farm
animal species produces milk similar to that of the mare,
finding a suitable formula is more complicated than just
substituting cow's milk, for example, for mare's milk.
However, goat's milk has been used successfully to raise
orphan foals. It is higher in fat and protein and lower in
carbohydrate than mare's milk, but foals drink it readily
and thrive on it, although some may experience diarrhea
(Koterba and Drummond, 1985). Interestingly, goat's milk is
similar to sow's milk, so perhaps this would further suggest
that piglets would be good models for foals.
Even though goat's milk and colostrum contain slightly
more protein and much more fat than that of the mare, foals
have been successfully raised at the veterinary teaching
hospital at the University of Florida on goat's milk


41
(Koterba and Drummond, 1985). Goat's milk is high in fat
and is similar to sow's milk (Glauser, 1966). Also, many
commercially available milk replacers for foals are fed at
twice the recommended concentration, which significantly
increases the fat intake (Naylor and Bell, 1985).
The compositions of colostrum and early lactation milk
from mares and sows are compared in Table 1.1. Ullrey et
al. (1966) and Pagan and Hintz (1985) reported data on
equine milk constituents, while Widdowson (1984) and
Widdowson (1985) reported on both sows and mares. While
similar in protein, sow's milk has higher total solids,
higher fat, and lower lactose than mare's milk.
Milk Production in Mares
Compared to other species, mare's milk is unusually
high in water. To compensate for this, foals must consume
large quantities of milk and have a rapid body water
turnover rate (the halflife of body water for foals under 1
wk old is 2.5 d). From birth to 11 d, foals will usually
drink about 16.2 kg of milk per day, ingesting about 422 g
protein and about 9830 kcals (Palmer, 1985).
When mares were fed iso-caloric diets that differed
only in the proportions of fiber and fat, milk compositions
did not differ (Pagan and Hintz, 1986). Mares fed diets
that contained 1.25 times the National Research Council


42
Table 1.1.
Comparison of colostrum and milk produced by
sows3 and mares .
Constituent
Mare
Sow
Matter
11.6
20.1
Dry Matter:
FAT
15.0
42.0
PROTEIN
22.8
29.0
CARBOHYDRATE
58.8
24.0
Colostrum Constituent Mare
Sow
(gm/100ml milk)
PROTEIN 25.2
FAT .7
LACTOSE 4.6
17.8
4.4
3.5
*Widdowson, 1984
Ullrey et al., 1966


43
(NRC) recommendations for energy requirements for lactation
produced greater volumes of more dilute milk than did mares
fed adequate calories. The growth rates of the foals were
not different for the two groups. The authors concluded
that feeding excess energy does not improve foal
performance, and results in obese mares.
Special Nutrient Requirements of Neonates
It is is important to determine the special nutrient
needs of neonates when developing a suitable enteral
formula. Premature neonates, for example will have
different nutrient requirements from healthy, full term
newborns (Koterba and Drummond, 1985). These authors
suggest that besides the major nutrients, requirements for
folate, Vitamin E, cysteine, calcium, and phosphorous
requirements for premature foals need to be evaluated. They
have found that stressed foals require about 120 kcal/kg/d
and may be expected to consume 20-28% of their body weight
(BW) in milk per day. They also recommend feeding an
elemental type of diet, because stressed newborns may not
tolerate milk-based diets which can cause bloat, colic, and
diarrhea. Elemental diets may prove to be a good
alternative to feeding cow's milk or soy-based formulas that
have been shown to have damaging effects on the intestinal
villi of infants (Weaver, 1986).


44
By examining prepartum mammary gland secretions, we may
be able to determine which nutrients are of special benefit
to premature neonates. In humans, the mammary gland
produces considerable amounts of medium chain fatty acids
(C8-C14) after parturition (Bitman et al., 1986). Linoleic
(18:2) also increases when compared to prepartum mammary
gland secretions. The secretion of several long chain
polyunsaturated fatty acids (PUSFA) decreases. Prepartum
secretions contained large amounts of C16 and C14 when
compared to normal milk. After parturition, the milk
contained increasing amounts of short and medium chain
length fatty acids (4:0, 6:0, 8:0, 10:0) as well as 18:0 and
18:1. Similar patterns of increased lipid synthesis and a
shift towards medium-chain fats after parturition were also
observed in cows. These authors suggest that the increase
in MCT in the milk may be of some advantage to preterm
neonates whose capacity for lipid digestion may not be
completely matured.


CHAPTER II
EVALUATION OF AN ELEMENTAL, ENTERAL DIET FOR NEONATES
USING PIGLETS AS A MODEL
Introduction
As recently as 20 yr ago, premature or sick human
infants were not expected to survive the neonatal period and
often were allowed to starve to death (Koterba et al.,
1985). Today, however, the advent of improved respiratory
therapy, refined surgical techniques, neonatal intensive
care facilities, and specially trained personnel gives the
very low birth weight or compromised infant a dramatically
improved survival rate. Once these infants could be kept
alive for longer periods of time, the question of nutrition
became important.
The field of equine neonatology, which has expanded
considerably over the past few years, has benefitted from
the advances made in human neonatology. Today, premature
and sick foals are treated successfully using much of the
same equipment and techniques developed for infants. Many
of these foals have gone on to achieve success on the race
track and in the show ring. Veterinarians are now faced
with the same questions faced by neonatologists treating
45


46
infants. Now that life threatening infections and
respiratory problems can be overcome (Webb et al., 1984;
Baker and Ames, 1987) what type of nutrition is appropriate
for these patients?
Ethical considerations make some types of research
impossible on human infants. Animal models have proven
useful in situations where invasive techniques or use of
radioisotopes is necessary. It was suggested that neonates
of different species may be more similar to each other than
they are to the adults of their species (Koterba et al.,
1985). Therefore, it is possible to use information gained
from work in one species for use in another. For example,
Rossdale (1987) used the human neonatology terms premature,
dysmature, and small for gestational age, to develop
guidelines for assessing readiness for birth in newborn
foals.
Although the elemental, enteral diets used in this
trial was designed originally for use in humans,
scientifically controlled trials on neonates were needed.
It was thought that this formula might be suitable for
premature or sick foals which were unable to drink mare's
milk. These compromised neonates may have poorly
functioning or damaged intestinal tracts.
Nutrients in enteral diets are delivered to the
digestive tract usually by mouth or by catheters inserted in
the stomach or small intestine. In contrast, parenteral


47
formulas are delivered intravenously. In an elemental diet,
individual nutrients are added in easily digestible forms,
in contrast to conventional, milk-based formulas. The base
formula tested in this study contained hydrolyzed whey
proteins rather than whole milk proteins. The formula was
lactose-free because, as stated previously, lactase may be
deficient when there is damage to the intestines. Milk-
based formulas contain long chain lipids almost exclusively,
but this formula included some medium chain fats. This was
because the digestion, absorption, and metabolism of long
chain fats may be inefficient in compromised neonates.
Piglets were used as a model for the preliminary
research in this trial. The information gained from these
trials would then be used as the basis for studies on foals
and infants. Colostrum-deprived piglets were used for
several reasons. If the piglets were allowed to nurse the
sow, it would not be possible to ensure that each piglet
received the same amount of colostrum. By hand-feeding the
piglets from birth, the amount and type of each nutrient
consumed can be controlled. Many sick neonates do not
receive colostrum shortly after birth and so are deprived of
its growth-promoting factors. The colostrum-deprived piglet
model permits the formula to be tested on stressed animals.
The restrictions and limitations of research projects with
foals and infants encourages the use of an appropriate
animal model for all preliminary work.


48
This study was part of a multi-project program to
develop an enteral diet for premature or sick foals.
Designing such a diet is a complex task requiring a series
of separate trials to determine the ideal formulation.
The goals of this specific study were:
1. to learn techniques and methods used in the investigation
of neonatal nutrition using a piglet model;
2. to determine the effect of an elemental, enteral diet on
the growth, intestinal development and general health of
colostrum-deprived neonatal piglets and, secondarily, to
learn to formulate diets with different lipid sources;
3. to evaluate and improve surgical and daily care
procedures for colostrum-deprived piglets hand-raised for 7
d; and,
4. to use the information obtained from the piglet study
towards possible development of a diet for neonatal foals.
Materials and Methods
Animals
This study was conducted using newborn Yorkshire x
Hampshire x Duroc crossbred piglets. The sows were bred and
cared for by the staff of the Reproductive Physiology Unit
under the direction of Dr. Fuller Bazer of the Animal
Science Department. Labor was induced 2 d prior to the
expected farrowing date to ensure that the birth would be
attended. The sow was injected with Prostaglandin F2a at


49
730 h on d 111 of gestation. Oxytocin was administered at
900 h on d 112. Farrowing usually occurred within several
hours after the oxytocin injection. All piglets were
removed from the sow immediately after birth, before
nursing. The umbilical cord was tied and cut. Identifying
markings, sex, weight, and general condition were recorded.
Only piglets greater than 750 g were admitted to the trial.
The protocol that subseguently was followed depended on the
treatment assignment of the litter.
Piglets from a total of 10 different litters were used
for the trial. Only one litter was on trial at any one time
due to time, labor, and facility constraints.
Baseline values were obtained from 18 piglets from two
litters removed to the animal facilities at the Food Science
and Human Nutrition Building and euthanized within a few
hours after birth. These non-fed piglets are referred to as
birth piglets (B0).
Normal values for sow-fed animals (SF) were obtained
from 22 piglets from two litters were returned to the sow
and allowed to nurse for 7 d, after which they were
euthanized. The sow and piglets were housed in an
environmentally controlled room. The sow was confined to a
farrowing stall of pipe construction. The piglets could
enter the stall easily under the bottom pipe or retreat out
of the stall to a corner of the room to avoid the sow.
Rubber floor mats were used to reduce conductive heat loss


50
and improve footing. Clean hay was provided for bedding in
a corner of the room where the piglets often slept. A heat
lamp provided additional warmth. The SF were monitored and
weighed daily.
Thirty-eight experimental piglets from six litters were
transported to the Piglet Neonatal Intensive Care Unit
(PNICU), an environmentally controlled room (37C, 70%
humidity) similar to hospital nurseries designed for human
neonates. These colostrum-deprived piglets were hand-raised
in individual plexiglass boxes (measuring 14" x 14" x 17")
in the PNICU for 7 d.
Surgery
All experimental piglets except those bottle-fed the
milk-based diet underwent surgery. Umbilical artery and
bladder catheters were inserted to permit collection of
blood and urine, respectively, during the trial. The
samples were used to evaluate the health of the piglets.
Gastric catheters were inserted to permit feeding by
constant infusion pump.
Development of anesthesia procedures. Piglets were
anesthetized for surgery with intramuscular injection (75
mg/kg BW) of ketamine HC1. Lidocaine was given
subcutaneously at the incision site. Once an intravenous
(i.v.) catheter was placed in the umbilical artery, an
Acepromazine drip was started at 60 drops/min. Full


51
recovery from this anesthesia was slow, delaying the
initiation of enteral feeding.
In an effort to decrease recovery time, the last two
litters on the trial were given the inhalant isoflurane by
mask as the sole anesthetic. Schieber et al. (1986) found
that although isoflurane caused a reduction in blood
pressure, peripheral resistance was reduced, preventing a
decrease in cardiac output. These workers suggest that
isoflurane, well tolerated by newborn piglets, has a clear
advantage over the use of halothane. The piglets in the
present study tolerated isoflurane well, recovering quickly
(some were quite active within minutes of removal of the
inhalant).
Catheter placement. Each piglet was fitted with an
umbilical artery catheter for administration of i.v. fluids
(.45% saline and 5% dextrose) during and after surgery, and
for collection of blood on d 3 and d 5 of the trial. The
piglets received the i.v. fluids until they were fully
recovered from the anesthetic and enteral feeding was
initiated, at least 12 h post-surgically.
Piglets also were fitted with bladder catheters for
urine collection. The ureters were left open to permit
normal urination.
The introduction of gastric catheters and the use of
continuous infusion pumps allowed enteral feeding to be


52
All catheters were externalized through a single
abdominal incision.
Post-operative care. After surgery, the piglets were
returned to their plexiglass boxes in the PNICU and observed
closely. A chart was kept beside each box for recording
data from individual piglets. Respiration, temperature,
color, activity level, and urine output, were recorded every
30 minutes until the piglets had recovered fully.
Diets
Hand-raised piglets were fed one of three enteral
formulations. A conventional, cow's milk-based formula was
bottle-fed to a total of 12 piglets from two litters. These
piglets, designated Group C, did not undergo surgery. In
addition, four piglets were fed this formula via gastric
catheter. Previous trials had shown that the growth rate of
piglets fed this formula was similar to the rate of sow-fed
piglets.
The two other diets used in this study were based on
the liquid elemental formula Peptamen (Carnation). All
piglets fed diets containing the elemetal formula were
designated Group E. Hydrolyzed whey proteins provide amino
acids and malto-dextrin is the carbohydrate source. The
formula was fed in two different, isocaloric forms based on
the predominant fat source. These were designated as MCT
(medium chain triglyceride) or LCT (long chain triglyceride)


53
the predominant fat source. These were designated as MCT
(medium chain triglyceride) or LCT (long chain triglyceride)
formulas. Because Peptamen is much higher in carbohydrate
than sow's milk, it was diluted to one-half concentration
before use. However, this necessitated the addition of whey
(27.4 g/liter for the MCT formula and 35 g/liter for the LCT
formula), calcium citrate (4.38 g/liter), and calcium
phosphate (5.77 g/liter) to obtain a composition similar to
sow's milk. The wrong type of whey mistakenly was added to
the formulas fed to one of the litters, invalidating the
data obtained from these pigs. The diets resembled sow's
milk in the amount of fat (60 g), protein (75 g), and
carbohydrate (63.5 g) provided per liter. Carnitine was
added at 486 il/liter of formula. The MCT formula was
obtained by adding 15 g medium chain triglyceride oil (MCT
Oil, Mead Johnson and Co., Evansville) and 6 g sunflower oil
(SFO) to 1 liter of Peptamen. All of the lipids in SFO are
long chain fats. The final MCT:LCT ratio was 70:30. The
LCT formula required the addition of .8 g MCT oil and 51.8 g
SFO to 1 liter of low-fat (LF) Peptamen. This resulted in a
90:10 LCT:MCT ratio. The composition of sow's milk and the
dietary treatments used in this trial are presented in
Tables 2.1-2.2.


Table 2.1 Composition of dietary treatments, base formulas, and sow's milk
on a per liter basis.
Diet
Composition
Protein
Fat
Carbohydrate
Calcium
Phosphorus
g
kcal
g
kcal
g kcal
g
g
Sow's milk
72
288
61
549
48 192
2.1
1.9
Peptamen
(U-5122-1)a
40
160
39
351
127 508
VO

o
in

o
Low Fat
Peptamen
(U-5216)b
47.6
190
7.4
67
186 744
0.6
0.5
MCT Formula
75
300
60
540
63.5 254.5
2.5
2.0
LCT Formula
75
300
60
540
93 372
2.5
2.0
Base formula
used
for MCT
diet.
DBase formula
used
for LCT
diet.


55
Table 2.2 Composition of sow's milk (Pond and Houpt, 1978)
and Peptamen (Carnation)
on a per liter basis.
Nutrient
Sow's Milk
Peptamen
Protein, g
72
40
Fat, g
61
39
Carbohydrate, g
48
127
Folic Acid, g
3900
4000
Thiamin, g
650
1500
Riboflavin, g
1370 -
1700
8200
Niacin, mg
4.3 9.0
20,000
B6' 9
200
3000
-6
B12, g
1.4
6 X 10
Biotin, g
14
.0003
Pantothenic
Acid, mg
4
10
Vitamin A, IU
500 8500
3750
Vitamin D, IU
100
200
Vitamin E, IU
1.4
20
Vitamin C, mg
146
100
Calcium, mg
2100
600
Phosphorus, mg
1000 -
500
1900
Potassium, mg
1000
1250
Sodium, mg
340
500
Magnesium, mg
200
300
Chloride, mg
1000
1000
Iron, mg
1.33
9
Zinc, mg
4.94
10
Manganese, mg
2.0
Copper, mg
1
Iodine, mg
.075
Energy, kcal
1030
1000


56
Development, of Feeding Regimen
The elemental diets were fed by bottle to the first
litter of piglets that received these formulas. Subsequent
piglets fed these diets were fed via gastric catheters. The
formulas were fed to the bottle-fed piglets at the rate of
15 ml/h on d 1 and 35 ml every 2 h on d 2. The rate was to
be increased by 5 ml/feeding on each successive day,
remaining at 50 ml/2h on d 6 and 7. Because the piglets in
this litter suffered from severe, watery diarrhea, the
formula was changed to Peptamen alone, at two-thirds
strength on d 5. Subsequent litters of Group E piglets, fed
via gastric infusion, received 10 ml h-1 kg BW-1 on d
1. On each successive day, the amount fed to healthy
piglets was increased by 3 h-1 kg BW-1. The flow rate
was not increased for piglets that were not thriving. The
last two litters were fed the MCT and LCT diets at half
strength. This decreased the production of watery feces.
With ketamine anesthesia, long post-surgery recovery
times delayed oral feeding. These piglets were often slow
to show normal sucking and swallowing behavior after being
anesthetized. Placement of a gastric catheter during
surgery allowed commencement of enteral feeding much sooner
than was possible with bottle-feeding. Enteral nutrition
was initiated on an individual basis when the piglet
appeared to be awake and kicking motions of the hindlegs
were observed. Usually, these animals remained on i.v.


57
fluids post-surgically for about 8 to 12 h before receiving
enteral nutrition.
To determine the effect of PNICU conditions on piglets,
the Group C controls were bottle-fed a milk-based formula
designed to be similar to the composition of sow milk. To
determine the effect of surgery on PNICU piglets, four
animals (surgery controls) were fed the milk-based diet via
gastric catheters using a continuous infusion pump. The use
of infusion pumps reduced the variation in post-absorptive
state that might occur when the animals are euthanized at
different times.
In summary, a total of 13 Group E piglets were fed the
elemental diets. Four piglets were bottle-fed, while the
rest received enteral nutrition via gastric catheter. Two
piglets were fed the base formula, Peptamen, with no
additions, via gastric catheter, to determine if the
additions of calcium and phosphorus were contributing to the
diarrhea. Of the 16 piglets fed the milk-based control
formula, 12 were bottle-fed and four underwent surgery for
gastric catheter placement and were fed via constant
infusion pump.
Daily Care
The piglets were monitored continuously throughout the
trial. Urine was collected asceptically from the bladder
catheters every 30 min. At morning and evening "rounds" the


58
incision site was checked and swabbed with an iodine
solution, the venous catheter was flushed, bandages changed,
and temperature, respiration, and girth measurements were
recorded. When piglets were fed via gastric catheter, new
bags of formula were started after rounds. Piglets were
weighed during morning rounds. On d 3 and d 5 blood was
collected from the umbilical catheter.
Every effort was made to keep stress on the piglets to
a minimum. Each piglet was provided with a rubber baby-
bottle nipple taped to the inside of the box, a plastic
ball, and a soft cloth.
Tissue Collection
At sacrifice, the animals were anesthetized with Na
pentobarbital and then exsanguinated by cardiac puncture.
Organs were removed as quickly as possible, weighed, and
frozen for later analyses. The intestines were measured,
weighed, cleaned of contents, and then re-weighed. The
small intestine of each piglet was divided into three
segments of equal length, designated proximal (top)
intestine (TI), middle intestine (MI) and distal (bottom)
intestine (BI).


59
Analyses
Leukocyte (WBC) counts were performed using a Unopette
dilution chamber and a hemocytometer. Microhematocrit (Hct)
determinations performed shortly after blood was collected.
Hemoglobin (Hb) was determined using the modified
Drabkins method with a Beckman DU-7 spectrophotometer at a
wavelength of 540 nm.
The plasma glucose was assayed using Trinder reagent
(Sigma Chemical Co., St. Louis), based on the glucose
oxidase reaction. The OD was read with the Beckman DU-7
spectrophotometer at 505 nm.
Non-collagen protein in intestinal homogenates was
determined using the micro-Biuret method (Itzhaki and Gill,
1964) after precipitation with .2 N NaOH. The OD was
measured with a Gilford spectrophotometer at 310 nm.
The activities of the intestinal disaccharidases
maltase, lactase, and sucrase were determined using a
slightly modified technique by Dahlqvist (1964).
Preparation of the tissue included homogenization in a
glass-on-glass hand homogenizer with .05 M sodium phosphate
buffer in a 1:5 weight/volume ratio. Each tissue homogenate
was incubated with each substrate to determine the
micromoles of glucose produced by the disaccharidase enzyme
per microgram tissue, during a 1 h incubation. The OD due to
the glucose produced was read at 505 nm with the Beckman DU-
7 spectrophotometer.


60
Urine was collected from Group E piglets every 30 min
throughout the trial. All of the urine collected during a 6
h period was pooled, unless it was unusual in appearance.
In this case, the abnormal urine was placed in a separate
vial. The urine was kept on ice or refrigerated until
analysis.
The volume of urine collected as well as a description
of its general appearance were recorded for each piglet for
each 6 h period. Fresh urine was analyzed using Ames
Multistix SG reagent strips (Miles, Inc., Ames). These
strips provided a quick assessment of specific gravity, pH,
protein, glucose, ketones, bilirubin, blood, and urobiligen.
The remaining urine was frozen, pending further analysis.
The following equation was used to calculate the % BW
gained by the 7-d old piglets:
% BW wt gain = ((d 7 BW birth BW) -s- birth BW) 100.
The percent of body weight of each section of the small
intestine (TI, MI, and BI) was calculated using the
following equation: SI section wt(% of d 7 BW) =
(SI section wt t d 7 BW) 100. These results will be
referred to as %TIWT, %MIWT, and %BIWT.
Additional Information
Additional information and details of surgery, daily
care, necropsy procedures, and tissue analyses are described
by Baltzell (1988). The PhD dissertation involved the the


61
colostrum-deprived neonatal piglet model that was further
developed in this current work.
Statisical Analysis
Treatment means for SF and BO data only were obtained
by method of least squares ANOVA (Snedecor and Cochran,
1969) using the GLM procedures of the SAS Statistical
Software (Freund and Littell, 1981). The model included
treatment, litter(treatment), sex, sex*treatment, and
sex*litter(trt). Data from Group C and Group E treatments
were not analyzed because of the low numbers of animals in
each group.
Results
Numerical results are expressed as least squares means
SE, unless stated to be observed means.
General Observations
Sow-fed piglets. All of the SF piglets were active,
healthy and growing well at the end of the 7 d trial period.
They did not seem unduly stressed by the daily weighing
procedure.
PNICU piglets. Thirty-eight piglets were started on
trial. All of the Group C piglets werew healthy and active
through by d 7. Although the elemental diets were
formulated to be similar to sow milk, the Group E piglets
did not thrive. These piglets experienced diarrhea that may


62
were made for each litter. These changes reduced but did
not eliminate the diarrhea. The survival rate of Group E
piglets was low, as only nine of 20 survived for 7 days.
This was probably due, in part to diet and to dislodged or
faulty catheters. Piglets that had dislodged gastric or
bladder catheters were humanely euthanized. Because of the
low survival rate of the Group E piglets, only descriptive
results will be presented.
Occasionally, piglets were born with umbilical hernias.
This was not generally not a problem, except for one piglet,
who diet after a portion of the small intestine
strangulated. Blockages or strictures of the small
intestine were observed in four animals.
Behavior of PNICU piglets. The hours of care required
by the piglets allowed for close observation of their
behavior. Each piglet had a particular activity pattern
that was known to all the caretakers. Therefore, early
detection of problems was possible.
The non-surgery piglets, bottle-fed the milk based
diet, quickly became accustomed to the routine in the PNICU.
They were noisy and wanted attention from the caretakers.
Even during the first day, they began to recognize the pre
feeding routine. They had strong suckling reflexes. Play
was part of their daily activity, just as it was for the
sow-raised piglets. Although they were kept in individual
boxes, the plexiglass allowed them to see each other and the


63
boxes, the plexiglass allowed them to see each other and the
caretakers. Objects such as crumpled paper towels, plastic
balls, and pacifiers were used as toys.
The healthy surgery piglets also exhibited play and
attention-seeking behavior and did not seem hampered by the
i.v. or enteral infusion lines attached to them. Soft
cloths were used as bandages to cover the incision and keep
lines in place. These were ignored by the animals. Swivels
in the infusion lines kept the twisting and tangling to a
minimum. These lines were changed twice daily, when fresh
formula bags were set up.
Activity level (sleeping, lying down, alert, moving,
etc.) of each piglet was recorded on the individual charts
every half hour and changes from the previous record noted.
Nasal oxygen was available, if needed. An attempt was made
to cool (water or alcohol swabs) or warm (heating pad or
heat lamp) animals whose body temperatures were not in the
normal range.
Healthy animals generally produced an adequate amount
of relatively clear urine. Animals that had decreased urine
output or bloody urine generally had enlarged kidneys on
necropsy and often had fluid accumulation in the abdomen and
hindlegs.
In an effort to improve the survival and growth rate of
the piglets fed the elemental diets, modifications in the
formulas and daily intakes were made for each successive


64
litter. After the first litter, the calcium and phosphorus
additions were discontinued. The amount of whey added to
the base formulas was decreased. The formula strength and
flow rate during the first few days after surgery were
decreased. It was thought that these changes might decrease
the diarrhea, improve tolerance of the formula, and smooth
the transition to enteral nutrition after surgery.
Gross Observations at Necropsy
Sow-raised and BO piglets. There were no abnormalities
noted in the organs of any of the SF or the BO piglets. The
intestinal tissue of the BO piglets was very thin compared
with that of the SF and could be easily torn when separating
the mesentery from the small intestine.
The stomachs of the BO piglets essentially were empty,
while the stomachs of SF piglets contained milk curd. The
proximal intestinal contents of the SF piglets were very
liquid and yellowish. The contents became thicker in the
more distal areas. The cecal contents were pasty and darker
yellow. Large intestine contents were very thick and became
darker in the more distal regions. The distal colon
contained formed fecal pellets, dark in color.
Post-surgical, formula-fed piglets. In all cases, the
positions of the catheters were verified on necropsy. No
abnormalities, other than occasional adhesions, were noted


65
with the catheters. The lining of the stomachs and bladders
did not show irritation from the catheters.
Some piglets exhibited respiratory distress and had
reddish, congested lungs at necropsy. Some of these animals
had displayed swollen shoulders, hindlegs, and abdomens with
bluish blotches seen beneath the skin. While cause of these
blotches is unknown, it may have been due to caretakers
holding the piglets too tightly during rounds.
Many animals with bladder catheters had enlarged
kidneys. In some cases, the ureters also were enlarged.
The livers of all of the piglets appeared normal, but
several piglets had gall bladders that contained a dark,
thick material. This was not seen in the SF or BO groups.
Non-surqical, formula-fed piglets. The Group C
piglets, fed the conventional, milk-based diet, were healthy
and active at the end of the 7 d. The organs of these
piglets appeared normal at necropsy.
Some piglets bottle-fed the elemental formulas had
slightly reddish, congested lungs.
Weight Gain
The means reported in this section are observed means
SE.
Sow-fed litters. The sow-fed piglets (SF) increased
their birth weight by 65.75%. One litter (n=13) gained an
average of 74.49% 4.24, with a minimum of 50.76% and a


66
maximum of 115.18%. The other litter (n=9) gained an
average of 53.13% 5.90. The least amount gained was
27.58%, and the most 79.16%.
Formula-fed piglets. The birth weight of Group C
piglets (n=12) increased by an average of 78.56% during the
week long trial. The smallest gain was 54.40%, while the
largest was 96.41%. The surviving surgery controls, fed the
milk-based diet via gastric catheters (n=2), gained only
29.75%.
One piglet fed one of the MCT-based diets gained 5.63%
of its birth weight and one piglet increased its weight by
21.91%. The other piglets experienced losses of 6.89%,
11.09%, and 8.5%. The MCT-fed piglets were all very thin by
the end of 7 d. None of the four surviving piglets fed
one of LCT-based lost weight, but they gained very little.
The average gain was 11.99%. The two bottle-fed piglets
gained 6.65% and 1.85% of their birth weight. The other
two, fed by gastric infusion, gained 35.29% and 4.18%. All
but two Group E piglets weighed greater than 1 kg at birth,
and these exceptions were over 900 g.
Small Intestine Length
The total length of the small intestine (SI) was
measured at necropsy. The length of the SI in the B0
piglets was used as a reference baseline value.


67
The least squares means (LS means) for one litter of BO
and one litter of SF piglets were 284.39 cm 11.27 and
371.73 cm 9.34, respectively.
Blood Glucose
Blood glucose may be expected to vary depending on an
animal's nutritional status, health, and physical condition.
The use of infusion pumps may have reduced variations due to
differences in times after feeding at death. This was
difficult to control with bottle feedings due to time and
labor constraints. The results in this section are reported
as observed means unless stated otherwise.
The LS means of blood glucose concentration of two
litters of SF piglets and one litter of BO piglets were
13.37 mM 1.81 and 6.55 mM 2.03, respectively.
Small Intestine Weight
The %TIWT, %MIWT, %BIWT observed for BO piglets, and
sow-fed piglets are presented in Table 2.3. The BO mean was
obtained from data from one litter and the SF mean was
obtained from two litters.
Disaccharidases in the Small Intestine
Results of the disaccharidase analyses are calculated
based on the amount of glucose produced by 1 g of tissue as
a result of substrate hydrolysis occurring during a 1 h


68
incubation period. The means for the enzyme activities for
SF and BO piglets are presented in Tables 2.4-2.6. The
results for the SF and BO piglets are from one litter of
each group. Therefore, the standard errors in the tables
also include litter effects.
Group C piglets (n=12), fed a milk-based formula, and
the Group E piglets had very low intestinal sucrase
activities. In tissue samples from several Group E piglets
no sucrase activity was detected.
Maltase activity was detected in Group E piglet
intestines, as well as in tissue obtained from Group C
piglets even though the Group C piglets had not ingested
maltose. Similarly, lactase activity was detected in the
Group E piglets that had never ingested lactose.


69
Table 2.3. Least squares means for the proximal, middle, and
distal sections of the SI, expressed as a
proportion of body weight.
Treatment
%TIWT
SE
%MIWT
SE
%BIWT
SE
Sow-fed (SF)
.83
.03
.84
. 03
.87
. 03
Birth (BO)
CO
in

. 04
. 63
. 03
. 62
. 04


Table 2.4. Least squares means of sucrase activity in proximal, middle, and distal
sections of the small intestine of 7-day old sow-fed piglets and
piglets killed at birth.
Sucrase Activity
nmol glucose produced/h/g wet tissue
TI
MI
BI
Treatment
Mean
SE
Mean SE
Mean
SE
BO
1.2
. 85
<.l .94
1.2
.70
SF
6.6
1.18
3.2 1.31
. 6
.98


Table 2.5. Least squares means of maltase activity in proximal, middle, and distal
sections of the small intestine of 7-day old sow-fed piglets and
piglets killed at birth.
Maltase Activity
/nmol glucose produced/h/g wet tissue
TI
MI
BI
Treatment
Mean
SE
Mean
SE
Mean
SE
BO
14.1
3.84
14.2
3.74
11.7
.93
SF
44.2
4.22
46.7
5.19
23.2
1.29


Table 2.6. Least squares means of lactase activity in proximal, middle, and distal
sections of the small intestine of 7-day old sow-fed piglets and
piglets killed at birth.
Lactase Activity
Mmol glucose produced/hour/gram wet tissue
TI
MI
BI
Treatment
Mean
SE
Mean
SE
Mean
SE
BO
45.1
8.89
217.2
24.23
82.3
8.07
SF
34.4
12.35
171.3
33.65
74.4
11.21


73
Complete Blood Counts
The complete blood counts (CBC), consisting of Hb, Hct,
and white blood cell counts, were performed to determine if
the blood components measured fell within published normal
ranges for piglets of this age. The CBCs provided a method
of monitoring the general health of the piglet during and
after 7 d on the trial. However, it was difficult to
collect blood from the umbilical artery catheter on d 3 and
5 of the trial as the catheters often became blocked and
only a very small amount of blood, if any, could be
withdrawn. Not enough data was collected on these days to
provide meaningful results. Data from SF and BO piglets is
presented in Table 2.7.


74
Table 2.7. Results of CBCs from blood collected at
necropsy via cardiac puncture from sow-fed
piglets and piglets killed at birth.
Treatment
na
Hb
(g/di)
Hct
(%)
WBC
(cells/ml3)
SF
22
5.6-10.5
18.7-35.3
7000-18,700
BO
9
3.3-20.5
10.3-42.2
4050-10,100
7_d h
sow-fed
31
3.6-5.2
24.1-34.9
8900-12,700
i-d b
sow-fed
70
12-12.7
39.6-43.5
6270-17,600
anumber of
samples
obtained
DSchmidt and Tumbleson, 1986
cd 28 data


75
Urinalysis
After surgery to introduce a bladder catheter, the
piglets routinely had some amount of blood in the urine.
Piglets with low urine outputs tended to have urine than was
visibly bloody, while piglets with substantial urine output
tended to have clear, yellow urine that was found to contain
blood only by chemstrip analysis.
The urine routinely contained varying amounts of
protein. This also may be expected after surgery. The pH
of the urine varied among piglets, but an individual piglet
tended to have urine in a small pH range throughout the
trial. The apparent health of the piglets did not seem
related to the changes in urine pH or specific gravity.
Discussion
The purpose of the work using piglets was to learn to
develop techniques and procedures to be used in a trial
designed to evaluate an elemental, enteral diet for sick or
premature neonates. To achieve this, two elemental formulas
and one milk-based formula were fed to the neonatal
colostrum-deprived piglets for 7 d. The elemental formulas
differed in the amount of fat provided by LCT and MCT
lipids. The elemental diets contained malto-dextrin as the
carbohydrate source, rather than lactose because compromised
neonates may have reduced lactase activity.


76
The animals in the first litter were bottle-fed the
elemental diets. None of them appeared to thrive during the
7 d study period, probably due to diarrhea and respiratory
problems. All subsequent Group E piglets were fed via
gastric catheters.
The diets were modified for each litter in an attempt
to develop a formulation suitable for the piglets. Changes
in the diet and flow rate reduced but did not eliminate the
diarrhea. Some workers had suggested that maltase may be
low in neonatal piglets. Therefore, it was likely that
undigested maltose reaching the large intestine may have
caused the diarrhea. This led to the investigation of the
carbohydrate source, maltose.
Disaccharidases in Neonatal Piglets
Human infants have adequate maltase at birth to digest
maltose even when they are born prematurely (Paige and
Bayless, 1981). However, there are conflicting reports in
the literature as to when maltase becomes active in the
newborn piglet. Very low maltase activity activities have
been detected in the small intestine of piglets as early as
2 d of age (Dahqvist, 1961a), and the activity appears to
increase from d 3 to 10 (James et al., 1987). The maltose
fed to the Group E piglets was within the amount reported to
be digested by normal, sow-fed piglets, but it is possible


77
that these colostrum-deprived piglets lacked sufficient
total tract maltase activity to meet their energy needs on a
maltose-containing diet.
Investigation of the tissue disaccharidase activities
in the small intestine of the experimental piglets was
undertaken to determine the ability of newborn and 7-d old
piglets to digest sucrose, maltose, and lactose.
Lactase activity was measured for reference as this
enzyme should be very active during the first week after
birth. Sucrase also would be a useful reference because
only low activities have been reported in 1-wk old piglets
(Veum and Mateo, 1986) .
The large variation in disaccharidase activities
between piglets in this trial was also reported by othe
workers (Manners and Stevens, 1972). It is difficult to
interpret the data when the large variation is coupled with
low numbers of animals per treatment.
The lactase activity along the SI was similar for the
BO and SF piglets. This is to be expected, because mammals
must be prepared, at birth, to digest lactose. The Group E
piglets, although not thriving, did have lactase activity,
even though they never consumed lactose. The intestine
remained prepared for the ingestion of this sugar throughout
the week.
The BO piglets had low maltase activities, leading to
the conclusion that piglets may not be able to digest


78
maltose efficiently enough to meet their energy needs when
this sugar is the sole carbohydrate source. The sow-fed
piglets appeared to have a numerically higher maltase
activity than did the piglets killed at birth even though
the SF piglets had never ingested maltose. This suggested
that age is a factor in enzyme development. Birth and
feeding, especially of colostrum, have been shown to
stimulate enzyme and intestinal development (Widdowson,
1985).
Sucrase activities were very low, compared with the
other disaccharidases measured. Dahlqvist (1961a) found no
appreciable sucrase in 2-d old piglets, but Beilis (1957)
reported equal digestion of lactose and sucrose by d 3.
Manners and Stevens (1972) were barely able to detect
sucrase activity until the piglets were 1 wk old. James et
al. (1987) found that EGF increased sucrose in neonatal
piglets. EGF is secreted in colostrum. In the present
study, only the SF ingested colostrum and this may explain
the generally higher enzyme activities measured in this
group.
Growth Rate in Neonatal Piglets
The Group C controls grew at a rate comparable with,
that of the sow-fed piglets. While the hand-raised piglets
did not have to expend energy to maintain body temperature
or compete for food, they did not have the benefit of


79
colostrum or sow's milk. Even though the growth rate of
Group E piglets was poor, the growth rate of the Group C
piglets indicated that colostrum-deprived piglets can be
raised successfully raised for 7 d in an intensive care
facility. The unit used in this study was kept extremely
clean but not sterile.
Development of the Small Intestine in Neonatal Piglets
It was suggested that the ingestion of colostrum and
milk affect the growth and development of the small
intestine in newborn mammals (Widdowson, 1985). An ideal,
enteral formula should supply adequate nutrients and growth
factors, especially to the small intestinal tissue. The
piglets killed at birth had very small, fragile intestines
that could be distinguished easily from the intestines of
older, fed piglets. By 1 wk of age, the small intestines of
SF piglets were longer and heavier than the BO intestines.
Because these weights are on a wet tissue basis, it is not
known whether the heavier weights are due to fluid or dry
matter.
Blood glucose
Although blood glucose is regulated by insulin and
glucagon, it can be affected by nutritional state, general
health, and time after feeding. Piglets are born with low
energy stores that must be replenished shortly after birth.


80
Blood glucose was low in BO piglets. These piglets often
were not sacrificed for several hours after birth, so
partial depletion of their energy stores probably occurred.
Normal blood glucose for neonatal piglets has been
reported by Tumbleson and Schmidt (198 6) The values had
large standard errors, which is expected as glucose varies
with time after feeding. They reported mean values for
birth and 1-wk old piglets of 2.67 1.06 and 6.67 1.11
mM/L, respectively. The birth value is lower than the 6.63
1.54 mM measured in piglets killed shortly after birth in
this study. All of the Group E piglets had blood glucose
concentrations similar to the reported values for 1-wk old
piglets. However, the 1-wk old SF and Group C piglets had
concentrations averaging about 13 mM. Method of blood
collection, stress, nutrition, breed effects, and time of
sampling have been known to affect results (Tumbleson and
Schmidt, 1986) .
Hematology
Results of the CBCs were compared with values reported
by Schmidt (1986). Hemoglobin in neonatal piglets was
reported to be 12.5 .2 g/dl at birth, and 10.0 1.8 g/dl
by d 7. The Hct values at birth are 40.3 .7% and 29.5
.4% by d 7. The piglets in the present study had
considerable variation in the Hb and Hct values, but most
were within or close to the reported ranges. However, the


81
PNICU surgery controls (n=2) had low Hb and Hct values.
These piglets had been very weak and may have had problems
unrelated to the treatment or surgery.
At birth, piglets would be expected to have about 6000-
1700 WBC//U1, and 7000-10,000 WBC//il by d 28 (Schmidt, 1986).
Generally, the piglets that survived until d 7 had CBC
values that were close to the normals for their age. Many
of the Group E piglets had severe diarrhea but did not have
extremely high packed cell volumes, suggesting that the
formula flow rate was sufficient to prevent dehydration.
Summary of Piglet Trial
As a result of this study, some techniques and
procedures for performing trials on colostrum-deprived
piglets were refined. Although each successive study will
probably result in further improvements, the colostrum-
deprived piglet model currently can be used successfully to
investigate the nutritional needs of neonates.
The poor growth rates of the piglets fed the MCT and
LCT diets may have been due to the carbohydrate source. No
information regarding the benefits of the addition of MCT to
neonatal formulas could be obtained from this trial.
However, using an appropriate base formula should allow this
information to be obtained from future studies using the
piglet model.


82
deprived piglet model currently can be used successfully to
investigate the nutritional needs of neonates.
The poor growth rates of the piglets fed the MCT and
LCT diets may have been due to the carbohydrate source. No
information regarding the benefits of the addition of MCT to
neonatal formulas could be obtained from this trial.
However, using an appropriate base formula should allow this
information to be obtained from future studies using the
piglet model.
It is clear that only digestible carbohydrates should
be included in neonatal enteral formulas. However,
information on the digestibilities of various sugars may not
be available for each species. Development of an enteral
diet for foals based on piglet data was a goal of this
research. However, it was not known whether newborn foals
had sufficient maltase activity to adequately digest the
malto-dextrins in the base elemental diet. Therefore, an
oral disaccharide tolerance test was performed on neonatal
foals.


CHAPTER III
ORAL DISACCHARIDE TOLERANCE TEST
FOR NEONATAL FOALS
Introduction
Neonates that are sick, stressed, or born prematurely
may have poorly functioning digestive systems. The pattern
of small intestine disaccharidase development may not be the
same for these newborns as for healthy neonates (Rossi et
al., 1986). Lactase activity can be reduced greatly as a
result of prematurity or disease (Rossi et al., 1986;
Tzipori et al., 1984). Consequently, lactose may not be
suitable for use in enteral diets designed to support non-
healthy neonates. The use of other carbohydrate sources
needs to be considered carefully. Including monosaccharides
in enteral diets would increase the osmolarity of the
formula higher than neonatal intestines could handle well,
possibly causing diarrhea. Complex sugars must be cleaved
into smaller and smaller units, and finally from
disaccharides into single sugar molecules before absorption
can take place. It is not known if neonatal foals can
digest and absorb disaccharide sugars other than lactose.
Therefore, it is important to determine which enzymes are
likely to be active in the intestine of equine neonates.
83


84
Enzyme activity varies with age and species and can vary
among similar individuals (Paige and Bayless, 1981). There
is very little information in the literature about the
disaccharidase activity in the small intestine of foals,
although it was studied in neonatal humans, pigs, calves,
and rats (Rossi et al., 1986; Dollar and Porter, 1957; Code,
1968; Paige and Bayless, 1981).
Dollar and Porter (1957) concluded from their study of
newborn calves that enzyme activities determined from
mucosal cell extracts were correlated closely to the results
of an oral disaccharide tolerance test. Using a similar
procedure, Roberts et al. (1974) studied lactase, maltase,
and sucrase activities in horses of various ages by
homoginizing samples of intestinal mucosa. Small intestine
tissue was obtained from the horses slaughtered or
euthanized. However, the data does not show the pattern of
disaccharidase development during the critical first week
postpartum. Lactase was detected as early as 105 d
gestation. Peak activity was found in foals at birth and it
remained high for 4 mo. Sucrase was barely detected in
fetuses from 3 to 9 mo gestation, remaining low at birth.
Adult activities were attained by 7 mo of age. Similarly,
maltase was very low until the ninth month of gestation.
Activities at birth were 12 to 15% of the enzyme activities
of adult equines. The authors reported a gradual increase,
until adult activities were reached at 7 mo of age.


85
In a subsequent study, Roberts (1975) determined the
ability of horses to digest and absorb disaccharides using
an oral disaccharide tolerance test. These results agreed
with those of his earlier study. However, this study did
not include data on very early neonatal life.
A non-invasive method of estimating disaccharidase
activity, such as an oral tolerance test, would allow foals
to be studied at more than one point in time and would not
require the experimental animals to be euthanized.
The present study was done using healthy foals, nursing
their mothers. Although the results will not necessarily be
applicable to sick or premature foals, it may eliminate some
disaccharides as possible ingredients for enteral diets; if
healthy foals are unable to digest a particular
disaccharide, then it is unlikely that a stressed foal would
have that ability. The results of this study will provide
information on the digestive ability of newborn, healthy
foals non-invasively, and help determine possible sources of
carbohydrates for use in enteral formulas. Because the
animals do not need to be euthanized to obtain samples, each
foal can be used on different days to determine the changes
in enzyme activity during the first week of life. There can
be considerable variation in activities among animals
(Manners and Stevens, 1972) .
Selection of a carbohydrate source for inclusion in
elemental, enteral formulas will depend on the activity of


86
the necessary enzymes and the desired osmolarity of the
formula. Disaccharidases are important in the digestion of
all sugars larger than monosaccharides.
It was presumed that healthy foals would have adequate
lactase activity while relying on milk for nourishment. It
was also presumed that glucose would be easily absorbed by
these neonates. Therefore, the changes in plasma glucose
occurring after administration of glucose and lactose would
be used for comparison with test sugars. Maltose and
sucrose were chosen as test sugars. Maltose was included
because it was the carbohydrate source in the base formula
fed to the piglets. Sucrose was tested because it was
thought that this sugar would be poorly digested and would
provide a reference for a non-absorbed substrate.
The goals of this trial were:
1. to investigate the ability of newborn, healthy
foals to digest maltose, lactose, and sucrose during the
first 5 d postpartum;
2. to investigate the change in blood glucose in
healthy foals after a short fast and the effects of age on
the ability of the foals to maintain blood gluocse
concentration after a short fast; and,
3. to determine the changes in blood glucose in
healthy foals in response to an oral glucose challenge.


87
Materials and Methods
Preliminary Study
A preliminary study to determine the effect of nursing
on blood glucose was performed on a normal, healthy foal at
8 h postpartum. Care of the mare and foal during and after
birth followed standard procedures at the University of
Florida Horse Research Center.
At 8 h of age, the foal was fitted with a Teflon
jugular vein catheter (Quik-Cath, Travenol Laboratories,
Inc.), in an aseptic procedure, to allow periodic blood
sampling. To minimize stress to the foal, a small amount of
local anesthetic was injected at the catheter site. The
plastic wings of the catheter were fixed to the skin with a
superglue adhesive. This was easily removed with acetone at
the conclusion of each trial period. The catheter was
filled with sterile, heparinized saline between samplings.
The foal was then allowed to return to the mare and resume
normal activity. After the foal nursed the mare, the time
was noted and the foal was muzzled to prevent nursing.
Blood was drawn from the catheter at this time (time = 0).
Blood was then drawn every 30 min thereafter for 2 h. This
procedure was repeated on the same foal on d 5. This study
suggested that blood glucose reached a baseline
concentration after a 1 h fast. It was concluded that a 2 h
fast prior to administration of the treatment, and a 4 h
fasting collection period would be sufficient to monitor


88
changes in blood glucose after a test meal in the subsequent
experiment.
Animals and Design
Eleven Quarter Horse and two Thoroughbred foals were
used for the main trial. The foals remained with their dams
throughout the trial. Foals were weighed shortly after
birth and on d 5. After each foal was born, it was randomly
assigned, by sex, to one of four treatment groups until the
block was filled. This procedure was repeated for
subsequent foals, assigning foals of each sex to fill the
treatment blocks evenly.
On d 1, at 6 h postpartum, each foal was fitted with a
jugular vein catheter, as described above. Foals were
muzzled to prevent nursing. The foals were then fasted for
2 h to allow blood glucose concentrations to reach a
baseline level. Each animal was then given the appropriate
treatment consisting of oral administration of either
maltose, lactose, or sucrose at 1 g (5.56 mmol) per kg BW in
a 20% solution (w/v), or a glucose solution at .5g (2.78
mmol) per kg BW. These dosages were similar to dosages used
by Roberts (1975).
The solution was offered to the foal in a bottle fitted
with a suitable nipple. If the foal did not voluntarily
consume the solution, it was administered via dosing syringe


89
or via nasogastric tube. Blood was collected from the
catheter immediately before the solution was given
(time = 0), every 15 min for the first hour, and every 30
min thereafter for the second, third, and fourth h of the
trial period. Each foal received the same treatment when
the procedure was repeated on d 3 and d 5 postpartum.
Analyses
The blood was collected in tubes containing
ethylenediaminetetraacetic acid (EDTA), plasma removed after
centrifugation at 3500 g for 20 min and frozen. Plasma
glucose was determined using the Trinder peroxidase method
(Sigma Chemical Co.). Optical density was measured at 505
nm with a Beckman DU-20 spectrophotometer.
Statistical Analyses
Statistical analyses were performed by method of least
squares ANOVA with time as a class variable (Snedecor and
Cochran, 1969,) using the GLM procedures of SAS (Freund and
Littell, 1981). The complete mathematical model is given in
Table 3.1. The appropriate error term is shown for each
independent variable.


90
Table 3.1. Complete mathematical model for ANOVA of changes
in plasma glucose.
Source df
Treatment 3
Sex 1
Treatment*Sex 3
Foal(Treatment*Sex) 5
Day 2
Treatment*Day 6
Day*Sex 2
Treatment*Sex*Day 6
Day*Foal(Treatment*Sex) 10
Time 10
Time*Treatment 30
Time*Foal(Treatment*Sex) 50
Day*Time 20
Sex*Time 10
Day*Sex*Time 20
Treatment*Sex*Time 30
Day*Treatment*Time 60
Treatment*Day*Sex*Time 60
Error term
Foal(Treatment*Sex)
Foal(Treatment*Sex)
Foal(Treatment*Sex)
Remainder
Day*Foal(Treatment*Sex)
Day*Foal(Treatment*Sex)
Day*Foal(Treatment*Sex)
Day*Foal(Treatment*Sex)
Remainder
Time*Foal(Treatment*Sex)
Time*Foal(Treatment*Sex)
Remainder
Remainder
Remainder
Remainder
Remainder
Remainder
Remainder
Remainder
101


Full Text
LIPID AND CARBOHYDRATE SOURCES FOR ELEMENTAL,
ENTERAL DIETS FOR NEONATES
By
LORI P. RICE
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1989

This dissertation is dedicated to my father,
Bernard Rudin,
who passed away in November, 1987. His humor,
wisdom, support, and encouragement, unselfishly
given to all who knew him, inspired great deeds.
He believed that I could accomplish anything I
set out to do. All things were possible.

ACKNOWLEDGMENTS
Sincere gratitude and appreciation are extended to the
chairman of my supervisory committee, Dr. E. A. Ott. He
always had patience, tolerance, and good spirits, even
during difficult times (such as trying to catheterize an
uncooperative foal at 3 a.m.). His guidance and knowledge
were essential to the completion of this project.
A special thanks is given to Dr. Peggy Borum, who
introduced me to the challenging field of neonatal
nutrition. Her dedication and high research standards were
an inspiration to do the very best job possible.
Support, encouragement, and advice were liberally given to
me by the other members on my advisory committee: Dr. E.
Johnson, Dr. D. Beede, and Dr. S. Lieb.
I am extremely grateful for the assistance of Fred
Buhl, who solved all my computer problems, Dr. Wilcox, who
very patiently helped with the statistical analyses, Mel
Tooker, and the crew at the Horse Research Center (C. Roden,
farm manager, Rose, Cindy, Jim, and Sharon). Maria
Fernandez and Sandy Duyck performed miracles restraining
fiesty foals and provided vital feedback. The piglet
projects benefitted from the experienced help of Janet
iii

Baltzell, Mike Sullivan, Sergio Quintana, and Robin Adkins.
My friends, Ty and Scott McClung, have been especially
helpful and supportive during my entire program.
Unwavering support, love, and encouragement were given
to me by my family. My husband, Brett, allowed himself to
be dragged 1000 miles to sweltering Florida and put up with
a student for a wife for longer than he'd like to remember.
My mother, Ellie Rudin, my grandmother, Ida Rudin, my sister
and her husband, Gail and Chuck Willis, were always there
when I needed them.
IV

TABLE OF CONTENTS
Page
ACKNOWLE DGMENTS iii
ABSTRACT viii
CHAPTER
I REVIEW OF THE LITERATURE 1
Introduction 1
Formulating Enteral Diets for Neonatal
Foals 2
Foals as Research Animals 3
Piglets as Models in Foal Research 5
Digestive Problems of Neonates 7
Infectious Causes of Diarrhea 7
Non-Infectious Causes of Diarrhea 8
Lipid Metabolism in Neonates 9
Fat in the Neonatal Diet 9
Fatty Acid Digestion and Absorption 11
Metabolism of Medium Chain and Long
Chain Fats by Neonates 13
Disaccharidase Activity in Neonatal Small
Intestine 15
Introduction 15
Brush Border Disaccharidases 19
Developmental Patterns of Enzyme Activity
in Neonates 20
Distribution of Disaccharidases 25
Sucrase 26
Lactase 27
Maltase 28
Determining Disaccharidase Activity in
Experimental Animals 3 0
Responses of the Neonate to Stress 31
Failure to Acquire Passive Immunity 31
Cold Stress 33
Compromised Neonates 3 6
Hematology and Blood Chemistry in Neonates.... 37
Hematology 37
Chemistry 39
Milk Composition and Intake 4 0
v

Page
Milk Production in Mares 41
Special Nutrient Requirements of
Neonates 4 3
II EVALUATION OF AN ELEMENTAL, ENTERAL DIET FOR
NEONATES USING PIGLETS AS A MODEL 45
Introduction 45
Materials and Methods 48
Animals 48
Surgery 50
Diets 52
Development of Feeding Regimen 56
Daily Care 57
Tissue Collection 58
Analyses 59
Additional Information 60
Statistical Analysis 61
Results 61
General Observations 61
Gross Observations at Necropsy 64
Weight Gain 65
Small Intestine Length 66
Blood Glucose 67
Small Intestine Weight 67
Disaccharidases in the Small Intestine... 67
Complete Blood Counts 73
Urinalysis 75
Discussion 75
Disaccharidases in Neonatal Pigs 76
Growth Rate in Neonatal Pigs 79
Development of the Small Intestine in
Neonatal Pigs 79
Blood Glucose 80
Hematology 81
Summary of Piglet Trial 81
III FOAL ORAL DISACCHARIDE TOLERANCE TEST 8 3
Introduction 83
Materials and Methods 87
Preliminary Study 87
Animals and Design 88
Analyses 89
Statistical Analyses 89
Results 93
General Observations 93
vi

Page
Fasting Blood Glucose 94
Glucose Absorption Time Curves 94
Treatment Curves Pooled Over All Days.... 95
Comparison of Day-Treatment Curves 100
Discussion 108
Fasting Blood Glucose in Neonatal Foals.. 108
Treatment Curves Pooled Over All Days.... 110
Day 1 Treatment Curves 112
Day 3 Treatment Curves 112
Day 5 Treatment Curves 114
IV CONCLUSIONS 118
APPENDICES
A PIGLET AND FOAL DATA 122
B DISACCHARIDASE ASSAY PROCEDURE 129
LITERATURE CITED 13 7
BIOGRAPHICAL SKETCH 145
vii

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
LIPID AND CARBOHYDRATE SOURCES FOR ELEMENTAL,
ENTERAL DIETS FOR NEONATES
By
Lori P, Rice
December 1989
Chairman: Edgar A. Ott
Major Department: Animal Science
Colostrum-deprived piglets were fed either elemental,
enteral diets, containing hydrolyzed whey proteins and
malto-dextrins (Group E), or a conventional milk-based diet
(Group C). Two litters were euthanized at birth (BO) to
provide baseline values. Normal values were obtained from
two litters of sow-fed (SF) piglets.
Group E piglets were surgically fitted with gastric,
bladder, and umbilical artery catheters, and fed by bottle
or continuous infusion pump. These piglets experienced
diarrhea. Diet and flow rate modifications, made for each
litter, reduced but did not eliminate the diarrhea.
To investigate diaccharidase activity in the small
intestine (SI) of these piglets, tissue homogenates were
incubated with maltose, lactose or sucrose. Maltase
activity was low in the SI of BO piglets suggesting that the
malto-dextrin in the elemental diet was probably not well
viii

digested by Group E piglets. Undigested maltose reaching
the large intestine probably caused the diarrhea.
Maltase activity was detected in the SI of Group C and
SF piglets even though they had never ingested maltose.
Similarly, lactose activity was detected in the SI of Group
E piglets that had never ingested lactose. Sucrase activity
was very low in all of the piglets.
To investigate disaccharidase activity in neonatal
foals, an oral tolerance test was conducted on 13 foals.
Maltose, lactose, sucrose, and glucose solutions were
administered orally on d 1, 3, and 5 postpartum. Changes in
plasma glucose from venous blood were measured over time.
Large peaks in plasma glucose occurred shortly after dosing
with lactose and glucose on all days. This suggests that
glucose from both substrates was absorbed. No rise in
plasma glucose was detected after oral sucrose
administration, even on d 5. Maltose was not well digested
on d 1. However, some digestion may have occurred on d 3
and 5, as evidenced by small increases in plasma glucose
after dosing with the maltose solution.
The colostrum-deprived piglet can be used successfully
as an animal model for nutritional investigations. The
results of the piglet and foal trials suggest that maltose
and sucrose would not be suitable substrates for these
neonates in the first few days of life.
ix

CHAPTER I
REVIEW OF THE LITERATURE
Introduction
Advances in medicine have decreased greatly the
mortality rate of weak or sick neonatal humans and animals.
New techniques in neonatology, especially in the field of
respiratory therapy, have enabled veterinarians to save many
diseased or premature foals that otherwise would not have
survived. The value, whether monetary or sentimental, of
many horses today has resulted in a greater number of high
risk foals receiving intensive care. As a result of the
great strides made in saving these foals, a growing number
of veterinary hospitals are developing or expanding their
facilities for the care of neonatal foals.
Once conditions that immediately threaten the life of
the newborn are stabilized, the primary concern becomes
administration of nutrients in a form that can be readily
utilized. In many cases, the foal can not or should not
consume mare's milk. Some foals must be separated from
their dams while they are in an intensive care setting.
Sick or weak foals that can not stand to nurse must be hand-
fed an appropriate enteral or parenteral diet.
1

2
Many of the feeding programs used on neonatal foals were
originally developed for human infants in intensive care
nurseries. Unfortunately, due to the difficulties in
working with these neonates, many of the formulas and
supplements being fed have not been well tested on very
young or stressed foals (Koterba and Drummond, 1985).
Formulating Enteral Diets for Neonatal Foals
Before an appropriate diet for stressed foals can be
formulated, basic research is needed to provide an
understanding of their metabolism. Many questions about the
efficiency of utilization of different nutrients remain
unanswered.
Ensuring adequate caloric intake is of primary
importance. In the past, some foals were inadvertently
starved to death while being hand-fed because their energy
requirements were underestimated (Koterba and Drummond,
1985). There is disagreement in the literature about the
amount and type of enteral diet that will supply adequate
nutrients during the critical neonatal period (Naylor and
Bell, 1985) .
Stressed or sick foals often have poorly functioning
digestive systems and may not absorb nutrients or antibodies
well. Some of these foals may even need parenteral feeding
to increase the supply of nutrients available to the cells.
Young foals are especially susceptible to infection when

3
they do not get an adequate supply of colostral antibodies
from their mothers, or when they are premature or dysmature
at birth (Rossdale, 1987). It has been shown in horses and
in other species that the nutritional needs of premature and
sick neonates can be different from that of term, healthy
newborns. While their need for nutrients remains the same
or even higher than that for healthy foals, their appetites
may be poor (Koterba and Drummond, 1985). Therefore, the
conclusions drawn from studies done on normal foals would
not necessarily be applicable to the weak or sick patients
treated in veterinary clinics.
Foals as Research Animals
Many problems present themselves when trying to design
experiments involving a relatively large number of research
foals. The expense of buying and feeding broodmares is
great, and there is no assurance that each mare will produce
a foal every year. Horses have a long gestation and tend to
foal during the same time of year. This makes it difficult
to spread foaling out over a long enough period of time to
facilitate handling many foals.
The labor required for round-the-clock care of even a
single foal is extensive. Even very young foals are
relatively large and require more elaborate facilities than
do small laboratory animals. Also, large radioactive
carcasses resulting from studies involving labeled compounds

4
would be difficult to dispose of. These concerns tend to
limit the number of foals that can be successfully studied
each year. These problems can be overcome, in part, by
using an appropriate animal model for preliminary
investigations.
Using animals as experimental models for human studies
is common practice. Ethical considerations have led to
extensive use of animal models for research concerning human
infants. Although the rat is frequently used as a model for
adults, rat pups are not similar to infants or foals.
Newborn rats have intestines that are fragile and difficult
to work with, and the rats' stage of development and rate of
growth at birth are very different from newborn humans,
pigs, horses, and ruminants (Widdowson, 1984). Calves,
while similar to foals, would not offer many advantages as a
model for foals.
Piglets have been used successfully as models for
premature infants. It has been suggested that they are
closer to human newborns than any other animal (Glauser,
1966). One of the objectives of this research is to
determine the appropriateness of using piglets as models
for the neonatal foal.

5
Piglets as Models in Foal Research
One of the major limitations of any animal model is, of
course, that the model is not going to behave exactly like
the real subject. Therefore the model chosen should always
come as close as possible to resembling the animal of
interest. There are no references in the literature to
previous work comparing piglets and foals, but information
on each shows that these neonates share many common traits.
The pig and the foal both tend to nurse freguently during
the day for short periods of time (Pegorier et al., 1983).
They have similar digestive functions at this age, both
being adapted to a milk diet. Also, both species depend
heavily on the passive immunity provided by the mother's
colostrum, which must be ingested shortly after birth.
It has been shown by many workers that foals and
piglets who do not acquire colostral antibodies are at
severe risk from many diseases including pneumonia,
septacemia, arthritis, and infectious diarrhea (Rossdale and
Leadon, 1975; Naylor and Bell, 1985; Rossdale, 1985; Koterba
et al., 1984; Mouwen, 1971; Bergeland and Henry, 1982). The
colostrum from both sows and mares contains predominantly
IgG (Pegorier et al., 1983; Naylor, 1979); however, IgA is
present in greatest quantities in non-colostral mare's milk.
Good management and sanitary conditions can help save
hypoglobulinemic neonates, but the risk of infection is
still high.

6
It has been suggested that mare's milk is closer to
that of primates than any other animal, being high in
lactose and low in fat and protein (Weaver, 1986). If
mare's milk is similar to human milk, and piglets are used
to simulate the response of infants, then perhaps the theory
that foals can also be compared to piglets has some
validity.
The use of the pig as a model has both advantages and
limitations. Being able to do research on piglets that are
littermates and on pigs from different litters, but by the
same parents, allows experiments to be performed using large
numbers of animals with minimal genetic variation. Also,
many animals can be studied in a relatively short period of
time, as the gestation of the sow (approximately 114 d) is
much less than that of the mare (approximately 335 d). Sows
can be bred to farrow all year, while mares have a foaling
season of several months.

7
Digestive Problems of Neonates
After birth, mammalian neonates undergo a period of
rapid acclimation to extrauterine life. They must adapt to
using enterally rather than parenterally supplied nutrients.
The neonate is now a free-living organism and it must be
able to ingest and absorb nutrients from its environment.
In mammals, of course, this usually means learning to nurse
from its mother. Serious problems occur when the newborn
has difficulty suckling or has an incompetent digestive
tract. Immature or damaged intestinal tissues will not
function normally, therefore, it is necessary to consider
the nutritional needs of neonates with digestive problems
separately from those of healthy neonates.
Diarrhea is a potentially serious problem for neonates
and can occur for several reasons. The causes are usually
classified as infectious or non-infectious.
Infectious Causes of Diarrhea
Diarrhea in piglets and foals has been attributed to an
assortment of viruses and bacteria. Rotavirus, in
particular, seems to be present in over 30% of the cases,
while coronavirus and adenovirus are also common infectious
agents (Palmer, 1985; Bergeland and Henry, 1982). These
authors suggest that rotavirus acts by causing damage to the
epithelium of the small intestine and leaving the neonate
vulnerable to a secondary bacterial infection. The bacteria

8
usually cultured in cases of piglet diarrhea include
Salmonella, and E. coli, while the latter is not considered
a problem in foals. They also suggested that in foals,
bacterial diarrhea may be secondary to septacemia and
enteritis. Piglets may also be plagued by coccidiosis or
infestations of strongyles.
Because of the ever present threat of invasion by
infectious organisms, the area where newborns are housed
must be kept extremely clean. It has been suggested that
foals usually become infected through contact with adult,
asymptomatic shedders on the same farm. Treatment for
diarrhea in piglets and foals usually includes
administration of antibiotics to cause a change in gut
flora, supplemental vitamins to offset decreased gut
absorption and bacterial synthesis, and, in piglets,
antispasmodics (Mouwen, 1971; Palmer, 1985).
Non-Infectious Causes of Diarrhea
Stress of all kinds, including exposure to low ambient
temperatures, or malfunctioning of the small intestine can
predispose newborn animals to diarrhea (Bergeland and Henry,
1982) .
Many foals suffer from diarrhea on d 6 to 14
postpartum. Because this is also the time when the mare
returns to estrus after giving birth, it has been called
"foal heat" diarrhea. However, there was no difference in

9
the amount or severity of diarrhea in hand-fed or mare-fed
foals observed by Palmer (1985). Therefore, the theory that
hormonal changes in the mare are reflected in the milk is
unfounded. It is important, though, to monitor the foal
closely at this time, because the onset of infectious
diarrhea can mimic foal heat diarrhea.
Piglets may excrete thin, watery feces called "white
scours" when they are very young. They usually seem normal
in attitude, but may be less active. The feces, upon
analysis, usually contains greater than 50% fat, but it is
not known if this is the cause or effect of the condition.
One experimenter suggests that it is the presence of
abnormally structured gut villi that is responsible for the
diarrhea (Mouwen, 1971).
Lipid Metabolism in Neonates
Fat in the Neonatal Diet
The addition of fat to human and animal enteral
formulas has proven to be an effective means of increasing
caloric density in an easily digestible form without also
increasing osmolarity (Koterba and Drummond, 1985). By
increasing the caloric density, the volume of formula to be
fed can be kept low. Large fluid loads can be detrimental
to sick or premature newborn patients. Fats are also added
to enteral and parenteral solutions to provide essential

10
fatty acids and to provide a carrier for fat soluble
vitamins.
Interest in studying the type of fat most readily
utilized by neonates has increased because the composition
of fat in fetal pig tissues and in sow's milk and colostrum
can be influenced by the source of dietary lipids fed to the
sow. Therefore, it may be possible to manipulate the diet
of piglets to affect the compostion of body lipids. This is
important because there is evidence that piglets metabolize
fats differently according to chain length and degree of
saturation (Miller et al., 1971; Lloyd and Crampton, 1957).
This also appears to be true in human infants (Faber et al.,
1988; Bach and Babayan, 1982; Lammi-Keefe and Jensen, 1984).
The fat content of milk and the extent of body lipid
stores at birth vary with species. There is usually an
increase in body fat of young animals after nursing.
Neonates who are energy-deprived must depend on fat reserves
for metabolizable energy and for thermal regulation
(Pegorier et al., 1983).
It has been suggested that within 2 d of birth neonatal
piglets can utilize fat as efficiently as glucose and the
ability to use fat increases with age (Miller et al., 1971;
Wolfe et al., 1978). Piglets have low fat stores at birth
(approximately 1% of their body weight) that rapidly
increase to about 15% of their body weight by day 14.
Wolfe et al. (1977) fed isocaloric diets to colostrum-

11
deprived, newborn piglets. The diets differed only in the
proportion of calories provided by carbohydrate and
butterfat. The fatty acid pattern of the backfat of the
piglets resembled the fatty acid composition of the diet,
especially when high fat diets were fed. Lipogenic enzyme
activity decreased with higher fat diets, which the authors
suggest is due to the inhibiting effect of increased lipid
intake rather than to decreased carbohydrate intake. Wolfe
and coworkers in 1978 determined that on low fat diets,
newborn piglets will resort to de novo fat synthesis and
attain a level of fat deposition similar to piglets fed a
high fat diet.
Fatty Acid Digestion and Absorption
Researchers have investigated the digestion and
absorption of lipids in neonates of many species, but more
information on how different types of fats are utilized by
premature and compromised newborns is needed. Data on lipid
metabolism in foals are limited.
Of particular interest is the effect of chain length on
neonatal absorptive and metabolic processes. In the past,
infant formulas contained exclusively long chain
triglycerides (LCTs), but recent work suggests that
compromised neonates may benefit from the replacement of
some of the LCT with medium chain length fats. It has been
suggested that medium chain triglycerides (MCT), usually

12
saturated fats consisting of 6 to 12 carbons, are readily
hydrolyzed and absorbed by infants, rat pups, and piglets
(Newport et al.; 1979, Putet et al., 1987; Jandacek et al.,
1987).
Medium chain fatty acids (MCFA) from the diet are
transported directly to the liver via the portal vein
whereas long chain fatty acids (LCFA) travel through the
lymph and then through the general circulation before
reaching the liver (Newport et al., 1979). MCT being more
polar and therefore more water soluble than LCT, are more
easily attacked by pancreatic lipase and thus, are a
desirable alternative to LCT in cases of pancreatic
insufficiency or in other situations when LCT would be
poorly absorbed (Jandacek et al., 1987).
Specifically designed triglycerides have been used to
benefit from both MCT and LCT in a single lipid source. A
triglyceride having medium chain fatty acids at the one and
three positions and a long chain fatty acid at the two
position may be the solution. This takes advantage of the
pancreatic lipase specificity for cleavage at the one and
three sites, leaving a 2-monoglyceride. Even in cases where
LCT absorption is poor, the 2-monoglycerides seem to be
easily transported across the intestinal mucosa (Newport et
al., 1979), thereby avoiding essential fatty acid (EFA)
deficiency. Human milk triglycerides have a 16:0 at the 2

13
position, which is thought to enhance the absorption of
dietary fats (Lammi-Keefe and Jensen, 1984).
Metabolism of Medium Chain and Lona Chain Fats
bv Neonates
The different metabolic pathways available to MCFA and
LCFA affect the energy sources available to peripheral
tissues. Interest in substituting MCT for LCT as part or
all of the lipid content of enteral formulas stems from the
products of metabolism of MCT. Several groups of
investigators have begun feeding MCT to neonates in an
effort to determine its benefits and drawbacks.
There is evidence to suggest that infants, especially
preterm infants, do better on MCT-added formulas than on
cow's milk alone, even though MCT are not found in large
amounts in mature human milk (Putet et al., 1987). Lammi-
Keefe and Jensen (1984) found considerably more 10, 12, and
14 carbon fatty acids in the milk of mothers that had very
premature or premature babies when compared to the milk of
mothers with full-term babies. They speculate that this may
be of some special benefit to premature infants. Some
researchers advocate feeding MCT to infants that have
malabsorption of lipids (Bach and Babayan, 1982).
MCT and MCFA are not normally found in the blood even
when they are added to the diet. They are thought to be
rapidly metabolized by the liver and do not reach the

14
peripheral tissues intact. However, 15-20% of the fatty
acids found in the umbilical cord blood of infants have
eight carbons or less. MCT tend to increase carbohydrate
tolerance, insulin secretion, and decrease glucose output
from the liver when given orally (Bach and Babayan, 1982).
A study on neonatal puppies (Cotter et al., 1987)
suggests that insulin causes LCFA to be removed from the
blood for storage, while MCFA are removed for immediate use
as an energy substrate. The authors have shown that low
doses of MCT given intravenously (i.v.) are utilized more
rapidly than LCT. The calories are available faster because
the lipoprotein lipase (LPL) in the capillary walls quickly
releases the MCFA, which then bind to albumin and are
absorbed by peripheral tissues. In the liver, MCFA can
enter the mitochondria rapidly, in contrast to LCFA which
must be activated and be transported with the help of a
carnitine carrier. MCFA can be oxidized so quickly that the
Kreb's cycle is overwhelmed. When this happens, the 2-carbon
units are funneled into ketone bodies which then diffuse
into the blood. The authors suggest that MCT is a more
flexible energy source than LCT. They also observed an
increase in plasma clearance rate of long chain fats when
medium chain fatty acids were present.
In a similar study, piglets fed either a 25:75 or 50:50
mixture of dietary MCT and LCT had higher blood glucose
concentrations and increased insulin secretion than pigs fed

15
a diet in which the fat source was 100% LCT (Wolfe et al.,
1978) .
Newport et al. (1979) fed 2 day old neonatal pigs
various ratios of MCT and LCT until they were euthanized at
day 28. The pigs fed a 50:50 ration had similar growth
rates to pigs fed 100% LCT as a dietary fat source.
However, 90% MCT feeding resulted in a significantly slower
growth rate. The pigs fed 90% or 50% MCT had higher liver
weights but lower carcass weights than the 100% LCT group.
The addition of MCT resulted in lower plasma lipid and free
fatty acids (FFA), but higher body water, total body lipids,
and cholesterol. They found no increase in nitrogen
retention due to MCT feeding as had been suggested by
others.
MCT feeding is not without possible disadvantages.
Urinary output of dicarboxylic acids (DCA) may be seen.
These end products of omega oxidation of medium chain fatty
acids, such as seberic, sebacic, and adipic, are often
detected in the urine during fasting, during ketosis, or in
children with defects in fatty acid oxidation or carnitine
metabolism. Henderson and Dear (1986) fed preterm infants
averaging 29 wk gestational age either parenterally (no
medium chain fats included) or enterally. The enteral
formulas contained either a high amount of MCT (1.8g/100ml)
or a low amount (.44 g/100 ml). Some infants received
breast milk containing .15 g MCT/100 ml. The infants fed

16
the high MCT formula excreted larger amounts of DCA than
infants fed the other feeding regimens, although no other
symptoms were evident. It is unclear if this would be a
problem for healthy infants. The authors stated that MCT
feeding would be unadvisable in infants with 0-oxidation
defects and that more investigation is needed to determine
the effects, if any, of increased urinary excretion of DCA.
They did not comment on the possibility that DCA excretion
resulted from inefficient use of MCT by the infants.
Ketones from MCT provide a guick source of energy for
neonatal extrahepatic tissues. As their concentration in
the blood increases, utilization by extrahepatic tissues
increases. Therefore, they are a good substrate for energy
production in times of high energy needs, such as growth or
undernutrition (Bach and Babayan, 1982). Ketogenesis is
controlled by a feedback system: when the concentration of
ketones in the blood increases, negative feedback to the
liver decreases production (Robinson and Williamson, 1980).
Gentz et al. (1970) have shown moderate increases in
plasma ketones during starvation in newborn and 16 day old
pigs during times when fat utilization is limited. They
suggest that this may not be detrimental because 10 kg pigs
are not prone to ketosis during starvation. Pegorier et al.
(1983) question the extent of ketogenesis and 0-oxidation in
newborn pigs since they do not show hyperketonemia as do
rabbits, rats, guinea pigs, and humans fed high fat diets.

17
In summary, the MCT system has a fast turnover rate but
is easily overloaded. The presence of medium chain fats
stimulate the absorption of plasma FFA. The 2-carbon units
resulting from /3-oxidation of MCFA are ketogenic, thereby
providing the flexibility of an alternative energy source.
The LCT system handles large quantities well but has a slow
turnover rate. The resulting increase in plasma FFA may
lead to acidosis. Some LCT are needed in the diet to
prevent essential fatty acid deficiency. MCFA may improve
the utilization and absorption of LCFA when there is
impaired lipid digestion.
Disaccharidase Activity in Neonatal Small Intestine
Introduction
Milk is a complex mixture of nutrients, hormones, and
other substances designed to provide optimal nutrition for a
normal, healthy newborn. Consequently, it is a difficult
task to formulate an artificial diet that will allow the
hand-raised neonate to grow and develop normally. The
challenge is even greater when the neonate is sick, weak or
premature.
The developmental pattern of intestinal disaccharidases
has been used to assess the nutritional status of neonates.
It has been shown that improper nutrition results in a
malnourished intestinal tract that does not show the same
pattern of enzyme activity changes as a healthy neonate

18
nursing its mother (Rossi et al., 1986). These authors
experimentally malnourished rats adding extra pups to a
normal-sized litter. The pattern of disaccharidase activity
in the malnourished rat pups resembled that of lesser
developed pups. The maltase and sucrase activities were
lower, and the lactase activity higher than in the control
pups.
Sick neonates often lose the ability to digest lactose
(Tzipori et al., 1984) and in infants, prematurity may
result in low total intestinal lactase activity (Mayne et
al., 1986). Therefore, it is important to assess the value
of alternative carbohydrates when developing artificial
formulas . Dairy farmers are also interested in the use of
economical energy sources for use in calf milk replacers
(Dollar and Porter, 1957). The efficient utilization of any
energy source will depend on the activity of appropriate
intestinal enzymes for digestion and absorption. Digestion
of complex carbohydrates, such as corn starch, are cleaved
by amylases into smaller units, into maltose, and finally
into single glucose molecules that can be absorbed (Code,
1968). Therefore, starch can not be utilized unless there
is adequate amylase and maltase present. Roberts (1974)
detected only low levels of both heat-sensitive pancreatic
a-amylase and heat-stable glucoamylase in neonatal foals.
One way to determine the digestibility of various sugars for

19
use in neonatal formulas is to estimate the amount of
disaccharidase activity present in the intestine.
Brush Border Disaccharidases
Disaccharidases are synthesized in membrane-bound
polysomes within enterocytes of the small intestine. There
is evidence to show that they are present in an inactive
form in crypt cells, becoming activated when the cells have
migrated to the top of the villus. Active lactase enzyme
has only been found in the microvillus membrane on "mature"
enterocytes at the tip of the villi in neonatal rats
(Mackenzie, 1985). The enzymes are activated when proteases
cleave a polypeptide chain from the large pro-disaccharidase
molecule. Extracellular pancreatic proteases are
responsible for activating maltase and the
sucrase-isomaltase complex, while lactase is activated by
intracellular proteases. The disaccharidases are attached
to the brush border of the epithelial cells lining the
intestine. In the rat, the rate of turnover for these large
proteins is short, about 11.5 h. The enterocytes themselves
take only 4 to 5 d to migrate from the crypt to the tip of
the villus, where they are shed (Dahlqvist and Semenza,
1985). The rapid rate of cell division and protein
synthesis makes these cells and their attached enzymes very
susceptible to adverse conditions and poor nutrition.

20
Disaccharidase enzymes, primarily sucrase, maltase, and
lactase, tend to follow the same pattern of development in
most mammals, but the rate of development, pre- and post¬
partum, differs by species (Paige and Bayless, 1981). The
amount of each disaccharidase present in the intestine will
determine the carbohydrates that can be successfully
incorporated into milk replacer formulas. Milk contains
many substances that can affect the development of the
intestines. Therefore, artificial formulas must be able to
support the normal pattern of maturation seen in neonates
nursing their mother. Determination of enzyme activity will
allow assessment of the development and general health of
the intestine. In this way, the nutritional value of
artificial diets can be compared to that of milk and
colostrum.
Developmental Patterns of Enzyme Activity in Neonates
The relative progression of the increase and decrease
in disaccharidase enzymes is similar for many animals, but
the timetables are very different for each species. For
most mammals, lactase is high at birth, increasing during
the nursing period, and decreasing near weaning. Lactase,
which develops in human infants later in gestation than
sucrase and maltase, reaches a peak at 36 to 40 wk gestation
(Roggero et al., 1986). Therefore, very premature infants
will have a limited ability to digest this sugar. Birth and

21
feeding each contribute to a postnatal rise in lactase in
both term and preterm infants (Weaver et al., 1986).
Studies on neonates of several species indicate
that prematurity or damage to the intestines can greatly
reduce lactase activity. Mayne and coworkers (1986)
developed a technique for assessing disaccharidase activity
in living human infants. They demontrated a significant
correlation between the enzyme activities measured in
jejunal fluid with enzyme activities in adjacent mucosal
tissue. Using this method, they studied eight premature,
but appropriate for gestational age infants 17 times during
the first 3 wk after birth. The infants were fed
pasteurized breast milk via nasogastric tube. The lactase
activity in these infants at 1 wk of age was lower than the
activity present at 2 and 3 wk of age (P < .001). The 1 wk
old infants had enzyme activities in the normal range for
adults. This work agreed with other published reports
indicating that lactase activity increases greatly near the
end of gestation in humans. This results in a lowered
capacity of preterm infants to digest a lactose-
based diet.
Damage to the intestinal tissue may also result in
reduced lactase activity. Biopsies were obtained proximally
and distally to a complete or incomplete congenital blockage
of the small intestines of 12 newborn infants during surgery
(Serrano and Zetterstrom, 1987). The activities of all

22
disaccharidases in tissue samples proximal to the
obstruction site were reduced when compared with previously
published reference values. Lactase activity was more
markedly reduced than the activities of the other
disaccharidases. There was no difference in lactase
activities in tissues from the four infants with complete
blockages compared the eight infants with incomplete
blockages.
However, when tissue samples from sites distal to the
obstruction were compared, lactase activity was lower (P <
.03) in infants with complete blockage when compared with
infants with incomplete blockage. The authors suggest that
distension and damage to the small intestine proximal to the
blockages resulted in impaired development of brush border
enzymes, particularly lactase. Minimal passage of ingesta
through an incomplete blockage permitted greater development
of lactase activity than detected in tissue distal to a
complete blockage.
To investigate the effect of infectious pathogens on
disaccharidase activities in neonates, Tzipori et al. (1984)
inoculated seven 4 day old foals with various combinations
of Streptococcus durans, Staphylococcus epidermis, and
Escherichia coli isolated from feces of foals with naturally
occurring diarrhea. Three foals received saline
inoculations. Inoculations were adminstered via
nasogastric tube and were followed with 200 ml evaporated

23
cow's milk. The foals were necropsied 1 to 3 d later. The
seven foals inculated with pathogens had experienced
diarrhea. S. durans bacteria were found to be adhered to
the intestinal mucosa. This probably contributed to the
significantly lowered lactase activity measured in these
foals compared with the lactase activity of the control
foals. These authors then conducted a similar study on
newborn piglets, inoculating them with the same bacterial
strains isolated from foals. These piglets also experienced
diarrhea and reduced lactase activity compared with saline
inoculated controls.
Maltase and sucrase do not appear in any appreciable
amount until at least a few days after birth (Code, 1968;
Veum and Mateo, 1986; Manners and Stevens, 1972; Ahrene et
al., 1969). This is the case for rabbits, calves, pigs,
rats, dogs, and cats (Code, 1968; Paige and Bayless, 1981).
Guinea pigs, however, are very mature at birth, nurse
infrequently, and can digest maltose and sucrose immediately
(James et al., 1987). Human infants develop considerable
maltase and sucrase activity before birth (Paige and
Bayless, 1981).
It has been reported that newborn piglets can utilize
lactose and glucose equally well at birth (Dollar et al.,
1957; Ahrene et al., 1969), while sucrase and maltase
activities increase during the first few days of life (Veum
et al., 1986). Dahlqvist (1961a) reported high lactase

24
activity in the pig at birth and a rapid increase in maltase
by day 2, but he found no appreciable sucrase at this age.
In contrast, Beilis (1957) reported that by day 3, piglets
could digest sucrose, lactose, and glucose equally well.
James et al. (1987) found low levels of maltase and sucrase
in sow-fed piglets on d 1 to 4 of life, with levels
increasing from day 3 through day 10. Sucrase activity was
twice that of maltase. When they gave Epidermal Growth
Factor (EGF), which may be present in colostrum, to
hand-raised piglets, they found an increase in sucrase and
maltase in the middle and distal portions of the small
intestine. EGF had no effect on lactase.
Dairy calf producers have tried to formulate milk
replacers using economical energy sources other than
lactose. Dollar and Porter (1957) fed newborn calves skim
milk using starch as an energy source with and without
amylase. Calves under 3 wk of age did poorly on this diet.
The authors suggest that the calves were unable to digest
the dextrins and maltose resulting from amylase digestion.
They report only very low activities of maltase and amylase
in the newborn calf and no sucrase activity. They state
that calves under 4 wk old have only been shown to digest
lactose and glucose. They did not see an appreciable rise
in maltose digestion until the calves were 9 wk old.

25
Distribution of Disaccharidases
The distribution of disaccharidase activity in the
small intestine has been investigated in several species.
In calves, lactase activity is highest in the proximal small
intestine. With age, lactase activity is reduce, mostly in
the distal third (Huber et al., 1961). Manners and Stevens
(1972) studied enzyme distribution in the newborn pig.
While they found great variation from pig to pig, they were
able to draw some conclusions. Enzyme activities were
measured at various sites along the small intestine, with
the proximal and distal ends represented as the 0% and 100%
sites, respectively. Lactase activity was highest at the
20% site along the small intestine, decreasing to a low at
the 95% site. From birth to 1 wk old, much of the decrease
seen in lactase activity occurred in the proximal 50%, with
higher activities found in the more distal section. Sucrase
activity was only barely detectable until the pigs were 1 wk
old, with the highest values in the proximal two-thirds of
the small intestine.
In foals, lactase activity was reported to be highest
in the duodenum and upper jejunum, decreasing distally
(Roberts et al., 1974). Maltase activity was low in the
equine fetus, increasing slowly during the first month after
birth mostly in the proximal and middle segments of the
small intestine, then increasing markedly until the end of
the first year. Sucrase was very low at all sites along the

26
intestine in the newborn foal, increasing in the proximal
sections during the first year, with the pattern of
distribution parallel to that of maltase.
Postmortem studies of the entire length of small
intestine of preterm and near-term human infants provided
information on the development of disaccharidases (Raul,
1986). The relative distribution of sucrase and lactase was
similar for infants in both age groups throughout the
intestine. The greatest amount of lactase activity was
found in the jejunum and duodenum, with little found in the
distal small intestine. Term infants had five times greater
lactase activity than the preterms, with the largest
differences found in the proximal intestine. The greatest
amount of sucrase activity was found in the jejunum. The
term infants had more enzyme activity in all segments of the
intestine. Glucoamylase was present in the tissues of
preterm infants, increasing distally, as it does in adults.
The authors suggest that this enzyme, along with salivary
amylase (which, in preterm infants, is still functional in
the small intestine) enhances digestion of glucose polymers.
Sucrase
Veum and Mateo (1986) fed pigs a sucrose-added formula
but could not stimulate sucrase activity by day 7. No
appreciable amounts of the enzyme were detected until day
14. Manners and Stevens (1972) found that pigs fed an

27
artificial diet had higher sucrase activity than sow-fed
controls. This may suggest that diet influences the rate of
development of the intestine. Huber et al. (1961) reported
that even when calves were fed a sucrose-added diet, there
was no notable digestion of sucrose until day 44, in
contrast to their study with pigs, where sucrase digestion
was evident by day 10.
Very low sucrase activity was detected in the small
intestine of equine fetuses and newborn foals. The activity
increased slowly during the first months after birth,
reaching adult activities by 7 mo of age (Roberts, 1974).
Lactase
Humans are an unusual species. For some races, the
ability to digest lactose remains long after weaning, but
the percent of the population with adult lactase activity is
small (Dahlqvist and Semenza, 1985). In humans, rabbits,
and rats prolonged feeding or absence of feeding of lactose
does not affect the postnatal changes in lactase activity
(Paige and Bayless, 1981). In calves, up to 44 d of age, no
increase in lactase activity occurs as a result of adding
lactose to the diet (Huber et al., 1961).
In human infants, lactose hydrolysis by the
/3-galactosidase enzyme is affected by other mono- and
disaccharides that may be concurrently ingested (Paige and
Bayless, 1981).

28
Lactase activity in the neonatal piglet has been shown
to be high at birth, reaching a maximum at day 15, then
decreasing throughout the nursing period with low activities
still detectable in the adult (Paige and Bayless, 1981; Veum
and Mateo, 1986; Manners and Stevens, 1971; Dollar et al.,
1957; Dahlqvist, 1961b).
Lactase activity was detected in the mucosa of eguine
fetuses as early as 105 d of gestation, increasing after the
ninth month, reaching a maximum at birth and remaining high
for 4 mo (Roberts et al., 1974). The activity declined,
especially in the duodenum, until 1 yr of age.
Maltase
Several researchers have investigated maltose as an
energy source for neonates. Cunningham and Brisson (1957)
fed 1 day old piglets a purified casein-based diet with
either maltose or glucose as an energy source. In piglets 3
to 7 d old, greater than 95% of the maltose in the diet, and
greater than 99% of the glucose disappeared from the small
intestine. The glucose diet resulted in greater
digestibility for protein and carbohydrate than the maltose
diet (P < .01). When the entire small intestine of neonatal
piglets of various ages was ligated and injected with a 5%
maltose solution, .66 /¿mol of maltose was hydrolyzed and
absorbed per h per kg body weight. This increased to 1.05

29
jLiinol by day 6 and 7. The authors conclude that maltose
digestion during the first wk may only be borderline in
supporting the animal's energy needs, but after day 7
maltose is used just as efficiently as glucose. Huber et
al. (1961) could not increase the maltase activity of calves
by feeding diets of whole milk plus lactose, or sucrose and
starch for 4 4 d. They note that this is in contrast to
other workers who have reported an increase in maltase
activity beginning in the third or fourth wk after birth.
Using intestines obtained from slaughterhouses, Roberts
et al. (1974) detected low maltase activity in equine
fetuses during the first 3 mo of gestation. Activities
remained low until birth. Enzyme activity doubled during
the first month, with the greatest increase occurring in the
proximal small intestine. Adult activities were achieved by
7 mo of age. No data were given specifically for the early
neonatal period.

30
Determining Disaccharidase Activity in Experimental Animals
Intestinal disaccharidase activity can be determined by
homogenizing samples of tissue from segments along the small
intestine and incubating the homogenate with disaccharide
substrates. The glucose produced is a measure of enzyme
activity. Many researchers have used this method, as
modified from Dahlgvist (1964). The major disadvantage of
this method is that the experimental animal must be
euthanized. Economic constraints usually limit the number
of foals that can be euthanized, making it impossible to
conduct large scale equine trials of this type. Therefore,
it would be helpful to have an alternate method, at least
during preliminary trials. Studies on calves have shown
oral tolerance tests to correlate well with the results
obtained using tissue homogenates when comparing relative
enzyme activities (Dollar and Porter, 1957).
Roberts (1975) used an oral disaccharide tolerance test
to determine the digestibility of various sugars in adult
horses and ponies. Although foals were not included in the
study, Roberts suggested that an oral lactose tolerance test
would be a useful tool for determining digestive competence
in the sick or premature foal. Testing the digestibility of
other sugars (for possible inclusion in artificial formulas)
in this way would allow animals to be studied without
euthanasia.

31
Responses of the Neonate to Stress
Stress factors include a wide variety of situations
that can cause problems for neonates. Poor maternal
nutrition and dystocia are examples of stresses can occur
before or during birth, but this project will be concerned
with stresses that occur during the neonatal period.
Failure to Acquire Passive Immunity
In the horse and pig there is little transport of
immunoglobulins across the placenta. So, these species must
depend on colostral transfer of antibodies to protect them
from a wide variety of infectious organisms until their own
immune systems are competent. The risk is high that these
neonates will be severely stressed by infections if they are
deprived of colostrum or are unable to absorb antibodies
during the first day of life. This becomes even more
serious if the neonate is also exposed to other stresses in
its environment or is weak, sick, or immature at birth.
Many researchers have emphasized the importance of
colostrum ingestion for newborn foals and have suggested
reasons why foals may be hypogammaglobulinemic even after
they have nursed (Naylor and Bell, 1985; Naylor, 1979;
Rossdale, 1985). Their findings show that immunoglobulins
in the colostrum decline rapidly, becoming 15% of its
concentration at parturition in 4 to 8 h.

32
Mares may produce poor quality colostrum for several
reasons. Older broodmares may leak colostrum before
parturition and so lose the high antibody content necessary
to provide the foal with adequate protection. Some foals
are born prematurely, before the mammary gland has completed
concentrating the immunoglobulins. Foals must ingest
colostrum within 24 h of birth. After this time, the
digestive tract "closes" and will no longer absorb large
molecules. Closure is not delayed by food-withholding.
Foals may be deprived of colostrum if they are orphaned
or because of a problem with the mare or with the foal
itself that prevents it from nursing. Foals that need to be
hospitalized may be separated from their mothers. Frozen
colostrum may either be unavailable or unwise to feed,
depending on the condition of the foal. Colostrum-deprived
foals have been shown to remain almost completely
agammaglobulinemic for 2 wk, then begin rapid synthesis of
antibodies until they appear the same as colostrum-fed foals
by 5 wk of age (Naylor, 1979). These workers feel that over
50% of colostrum-deprived foals will develop an infection
resulting in septacemia, diarrhea, pneumonia,
omphalophlebitis, or arthritis within 1 wk of birth. They
suggest prophylactic treatment of 1.2 liters of plasma taken
from an adult horse, given intravenously over 2 h, to supply
an adequate amount of IgG.

33
Piglets are also likely to suffer similar diseases if
not provided with passive immunity from colostrum (Svendsen
and Bille, 1981; Bergeland and Henry, 1982) or kept in
isolated, aseptic conditions (Pond, 1978). In addition to
providing passive immunity to neonates, colostrum also
contains epidermal growth factors that promote mucosal
development in the gastrointestinal tract (Svendsen and
Bille, 1981). Unfortunately, it has been shown that
colostrum-deprived neonates fed either milk from other
species or soy-based formulas sustain damage to the
intestine, as the immune system responds to the foreign
proteins ingested (Weaver, 1986). The use of elemental
diets would prevent this adverse reaction as they contain
hydrolyzed rather than whole proteins.
Cold Stress
Exposure to a low ambient temperature shortly after
birth is a common problem for many neonates including
piglets, foals, and human infants. Management practices
used on many breeding farms encourage mares to begin cycling
and ovulating earlier in the year than they would in nature,
so foals are born more and more freguently in late
winter/early spring when temperatures are low. Therefore,
it is important to study the ability of neonates to regulate
their body temperature and the effects of cold stress on
their health and nutritional requirements.

34
Normal body temperature for foals is 38.05°C ±1, with
the lower boundary of the thermal neutral zone being 14°C
during the first 48 h postpartum. After that, the foal can
tolerate an ambient temperature as low as 8°C, provided he
has nursed and the environment is free of draught (Rossdale,
1985). Premature foals often have difficultly in
maintaining their body temperature, which may drop to 36.6°C
or even lower (Naylor, 1979). Foals lack significant
deposits of brown adipose tissue and must rely on shivering
for thermogenesis (Rossdale, 1985).
The normal body temperature of piglets is 39°C at
birth. The thermal neutral zone is very small (34-35°C)
(Stanton et al., 1973) and they have greater difficulty
maintaining their body temperature under cold stress than do
foals and calves (McCance and Widdowson, 1959).
Piglets are easily cold stressed for several reasons:
they are poorly insulated at birth, having almost no brown
adipose tissue and only 1% of their body weight (about 10 g)
is fat; they have a small body size with a large surface
area; their metabolic rate is low at birth; and they are
born with no protective hair or fur (McCance and Widdowson,
1959; Gentz et al., 1970; Curtis and Rogler, 1970; Stanton
et al., 1973). However, their ability to thermal regulate
does increase with age and is fairly well developed by 1 wk
of age (Gentz et al., 1970; Curtis and Rogler, 1970; Stanton
et al., 1973) .

35
Piglets will experience a 2 degree decrease in body
temperature, to 37°C, shortly after birth. Adverse effects
may be seen if the temperature reaches 35°C (Svendsen and
Bille, 1981).
An ambient temperature of 5°C is close to the cold
limit for newborn pigs. At this temperature, there should
be maximum utilization of their thermal regulatory
capabilities. Several investigators have exposed newborn
pigs to this temperature for short periods of time. Results
suggest that starving piglets undergoing cold stress
experience hypoglycemia and will not survive longer than 30
h unless fed (McCance and Widdowson, 1959). These workers
report that at birth, pigs are able to increase oxygen
consumption during cold stress whether they are fed or not,
whereas the newborn rat will not show an increase until it
is 5 d old. Rabbit young improve their ability to respond
during the first few days of life.
Carbohydrate is the main energy source of newborn
piglets, even though sow's milk contains 30-40% fat on a dry
matter basis (Allee et al., 1971) because fat utilization is
limited during the first few days postpartum, regardless of
the ambient temperature. However, cold exposure results in
rapid depletion of energy stores and death from hypothermia
unless the piglets are fed. It is estimated that the
carbohydrate and fat stores endogenous to the piglet would

36
yield about 72 h worth of energy with ambient temperatures
in the thermal neutral zone (Gentz et al., 1970).
Compromised Neonates
Being born premature or dysmature will compromise the
neonate's ability to adapt to extrauterine life. Dystocia
and neonatal maladjustment syndrome (where there is reduced
ability to suck, swallow, or move about) also predispose
newborns to septacemia and in the case of foals, septic
arthritis (Koterba et al., 1984). Premature foals usually
have abnormal blood gases and an impaired acid/base status
(Rossdale et al., 1987). They often are weak and take a
longer-than-normal time to stand and nurse. They commmonly
show hypoflexion of the metacarpal and metatarsal joints,
and a bright red tongue, but they may maintain normal
respiration and heart rate for the first 24 h before their
condition deteriorates (Rossdale, 1987).
Intestinal maturation will not be complete in preterm
neonates. Enterocyte function, villi development, and
enzyme activity are often different from that found in term
newborns, resulting in decreased digestibility of some
nutrients (Raul et al., 1986; Mayne et al., 1986; Mackenzie,
1985).

37
Hematology and Blood Chemistry of Neonates
Hematology
It has been suggested that evaluating the packed cell
volume (PCV), hemoglobin (Hb), red blood cell count (RBC),
white blood cell count (WBC), and differential leukocyte
count can be helpful in assessing the condition of neonates
and estimating the degree of maturity and viability of
neonates (Rossdale, 1985).
Becht and Semrad (1985) found that in foals, PCV, Hb,
and RBC counts peak at birth and then begin to decrease
within 12 h postpartum. They also noticed that mean
corpuscular volume (MCV) decreased slightly in the fetus
just before birth, then remained steady for the first 2 wk
of extrauterine life. They suggest that some of these
indices may be useful for establishing the maturity of the
foal at birth.
Several workers have reported lower leukocyte counts (4
x 10 /liter) for premature foals as compared to full term
Q
foals (6 x 10 /liter). Erythrocyte counts were also lower (6
x 1012 vs 12 x 1012/liter) as was the PCV (33 vs 45.6%).
Globulin (<10 gm/liter) and gammaglobulin (4 gm/liter) were
also lower than for normal foals (Jeffcott et al., 1982;
Kitchen and Rossdale, 1975; Rossdale, 1983; Becht and
Semrad, 1985).
Foals have an intact granulocytic system at birth. The
increase in WBC reported during the first day of life is

38
thought to be due to an increase in mature polymorphonuclear
neutrophils (PMN) and lymphocytes. The increase in PMN
continues during the first 3 days postpartum. During the
first wk, monocytes begin to appear. Basophils are not
normally seen in the neonatal period.
9
Premature foals tend to have lower WBC (4 x 10 /liter) ,
RBC (6 x 1012/liter), PCV (33%), globulin (<10gm/liter),
gammaglobulin (4gm/liter) and a narrower neutrophil to lymph
ratio (1.2:1) than normal foals. A severe leukopenia, left
shift, and the appearance of toxic neutrophils may indicate
sepsis (Becht and Semrad, 1985).
Jeffcott et al. (1982) and Rossdale (1983) found that
premature foals consistently show a lower than normal
neutrophil to lymphocyte ratio (N/L). Normal foals have a
ratio > 2:1 while premature foals show a reversed N/L ratio
of > 1:1. A dose of short acting exogenous ACTH will elict
a neutrophilic response in mature foals but not in immature
foals that lack normal adrenal function.
In piglets, RBC (from 6.18 x 1012/liter to 4.4 x
1012/liter), PVC (from 40 to30%) , and Hb (12.5 to 10.0g/dl)
are decrease during the first wk after birth. However, the
number of WBC tends to increase from about 6.2 to 17 x
3
10 //Lil. The proportion of PMN to lymphocytes percentages
shifts from about 38:60 to about 53:42. Granulocytes are
rarely seen during the first wk (Schmidt and Tumbleson,
1985) .

39
Chemistry
Various clinical chemistry analyses are useful for
evaluating the general health and nutritional status of
newborn animals. Both foals and piglets are susceptible to
hypoglycemia shortly after birth, especially if they are
weak, sick or stressed.
Normal blood glucose in the neonatal piglet is 60 to 80
mg/dl. However, it may be as high as 100 mg/dl immediately
after birth. The decrease may be due to limited glycogen
stores in the liver (Pond, 1978). Blood glucose values as
low as 48 mg/dl have been reported for newborn pigs,
increasing to 114 mg/dl by day 7.
Foals are susceptible to hypoglycemia shortly after
birth, especially if they are weak, sick or stressed. Foals
that have suckled during the first 2 h after birth were
reported to have the following blood glucose values: 95.5 +
17.44 mg/100 ml at birth, 83 ± 3.28 mg/100 ml 30 min
postpartum, and 131 + 12.4 by 14 h (Kitchen and Rossdale,
1975). Normal, healthy foals that have suckled have blood
glucose levels that are higher than adult levels for the
first 24 h and remain in the high normal adult range for the
first 30 d. Hypoglycemia may result in sick foals not
receiving sufficient nutrient intake, or from poor digestive
and absorptive function even with adequate intake (Becht and
Semrad, 1985).

40
Blood lipid components such as free fatty acids,
trigycerides, ketones, blood urea nitrogen (BUN), and
circulating liver enzymes are important parameters to
consider when evaluating the utilization of dietary fat and
nitrogen status in the neonate.
Milk Composition and Intake
Some researchers, using milk replacers currently
available for foals, have expressed dissatisfaction with the
foals' growth rate and general appearance. They report that
foals fed milk replacers according to directions usually
remain small (Naylor and Bell, 1985). Since no other farm
animal species produces milk similar to that of the mare,
finding a suitable formula is more complicated than just
substituting cow's milk, for example, for mare's milk.
However, goat's milk has been used successfully to raise
orphan foals. It is higher in fat and protein and lower in
carbohydrate than mare's milk, but foals drink it readily
and thrive on it, although some may experience diarrhea
(Koterba and Drummond, 1985). Interestingly, goat's milk is
similar to sow's milk, so perhaps this would further suggest
that piglets would be good models for foals.
Even though goat's milk and colostrum contain slightly
more protein and much more fat than that of the mare, foals
have been successfully raised at the veterinary teaching
hospital at the University of Florida on goat's milk

41
(Koterba and Drummond, 1985). Goat's milk is high in fat
and is similar to sow's milk (Glauser, 1966). Also, many
commercially available milk replacers for foals are fed at
twice the recommended concentration, which significantly
increases the fat intake (Naylor and Bell, 1985).
The compositions of colostrum and early lactation milk
from mares and sows are compared in Table 1.1. Ullrey et
al. (1966) and Pagan and Hintz (1985) reported data on
equine milk constituents, while Widdowson (1984) and
Widdowson (1985) reported on both sows and mares. While
similar in protein, sow's milk has higher total solids,
higher fat, and lower lactose than mare's milk.
Milk Production in Mares
Compared to other species, mare's milk is unusually
high in water. To compensate for this, foals must consume
large quantities of milk and have a rapid body water
turnover rate (the halflife of body water for foals under 1
wk old is 2.5 d). From birth to 11 d, foals will usually
drink about 16.2 kg of milk per day, ingesting about 422 g
protein and about 9830 kcals (Palmer, 1985).
When mares were fed iso-caloric diets that differed
only in the proportions of fiber and fat, milk compositions
did not differ (Pagan and Hintz, 1986). Mares fed diets
that contained 1.25 times the National Research Council

42
Table 1.1.
Comparison of colostrum and milk produced by
sows3 and mares .
Constituent
Mare
Sow
â–  Matter
11.6
20.1
Dry Matter:
FAT
15.0
42.0
PROTEIN
22.8
29.0
CARBOHYDRATE
58.8
24.0
Colostrum Constituent Mare
Sow
(gm/100ml milk)
PROTEIN 25.2
FAT .7
LACTOSE 4.6
17.8
4.4
3.5
3Widdowson, 1984
“Ullrey et al., 1966

43
(NRC) recommendations for energy requirements for lactation
produced greater volumes of more dilute milk than did mares
fed adequate calories. The growth rates of the foals were
not different for the two groups. The authors concluded
that feeding excess energy does not improve foal
performance, and results in obese mares.
Special Nutrient Requirements of Neonates
It is is important to determine the special nutrient
needs of neonates when developing a suitable enteral
formula. Premature neonates, for example will have
different nutrient requirements from healthy, full term
newborns (Koterba and Drummond, 1985). These authors
suggest that besides the major nutrients, requirements for
folate, Vitamin E, cysteine, calcium, and phosphorous
requirements for premature foals need to be evaluated. They
have found that stressed foals require about 120 kcal/kg/d
and may be expected to consume 20-28% of their body weight
(BW) in milk per day. They also recommend feeding an
elemental type of diet, because stressed newborns may not
tolerate milk-based diets which can cause bloat, colic, and
diarrhea. Elemental diets may prove to be a good
alternative to feeding cow's milk or soy-based formulas that
have been shown to have damaging effects on the intestinal
villi of infants (Weaver, 1986).

44
By examining prepartum mammary gland secretions, we may
be able to determine which nutrients are of special benefit
to premature neonates. In humans, the mammary gland
produces considerable amounts of medium chain fatty acids
(C8-C14) after parturition (Bitman et al., 1986). Linoleic
(18:2) also increases when compared to prepartum mammary
gland secretions. The secretion of several long chain
polyunsaturated fatty acids (PUSFA) decreases. Prepartum
secretions contained large amounts of C16 and C14 when
compared to normal milk. After parturition, the milk
contained increasing amounts of short and medium chain
length fatty acids (4:0, 6:0, 8:0, 10:0) as well as 18:0 and
18:1. Similar patterns of increased lipid synthesis and a
shift towards medium-chain fats after parturition were also
observed in cows. These authors suggest that the increase
in MCT in the milk may be of some advantage to preterm
neonates whose capacity for lipid digestion may not be
completely matured.

CHAPTER II
EVALUATION OF AN ELEMENTAL, ENTERAL DIET FOR NEONATES
USING PIGLETS AS A MODEL
Introduction
As recently as 20 yr ago, premature or sick human
infants were not expected to survive the neonatal period and
often were allowed to starve to death (Koterba et al.,
1985). Today, however, the advent of improved respiratory
therapy, refined surgical techniques, neonatal intensive
care facilities, and specially trained personnel gives the
very low birth weight or compromised infant a dramatically
improved survival rate. Once these infants could be kept
alive for longer periods of time, the question of nutrition
became important.
The field of equine neonatology, which has expanded
considerably over the past few years, has benefitted from
the advances made in human neonatology. Today, premature
and sick foals are treated successfully using much of the
same equipment and techniques developed for infants. Many
of these foals have gone on to achieve success on the race
track and in the show ring. Veterinarians are now faced
with the same questions faced by neonatologists treating
45

46
infants. Now that life threatening infections and
respiratory problems can be overcome (Webb et al., 1984;
Baker and Ames, 1987) what type of nutrition is appropriate
for these patients?
Ethical considerations make some types of research
impossible on human infants. Animal models have proven
useful in situations where invasive techniques or use of
radioisotopes is necessary. It was suggested that neonates
of different species may be more similar to each other than
they are to the adults of their species (Koterba et al.,
1985). Therefore, it is possible to use information gained
from work in one species for use in another. For example,
Rossdale (1987) used the human neonatology terms premature,
dysmature, and small for gestational age, to develop
guidelines for assessing readiness for birth in newborn
foals.
Although the elemental, enteral diets used in this
trial was designed originally for use in humans,
scientifically controlled trials on neonates were needed.
It was thought that this formula might be suitable for
premature or sick foals which were unable to drink mare's
milk. These compromised neonates may have poorly
functioning or damaged intestinal tracts.
Nutrients in enteral diets are delivered to the
digestive tract usually by mouth or by catheters inserted in
the stomach or small intestine. In contrast, parenteral

47
formulas are delivered intravenously. In an elemental diet,
individual nutrients are added in easily digestible forms,
in contrast to conventional, milk-based formulas. The base
formula tested in this study contained hydrolyzed whey
proteins rather than whole milk proteins. The formula was
lactose-free because, as stated previously, lactase may be
deficient when there is damage to the intestines. Milk-
based formulas contain long chain lipids almost exclusively,
but this formula included some medium chain fats. This was
because the digestion, absorption, and metabolism of long
chain fats may be inefficient in compromised neonates.
Piglets were used as a model for the preliminary
research in this trial. The information gained from these
trials would then be used as the basis for studies on foals
and infants. Colostrum-deprived piglets were used for
several reasons. If the piglets were allowed to nurse the
sow, it would not be possible to ensure that each piglet
received the same amount of colostrum. By hand-feeding the
piglets from birth, the amount and type of each nutrient
consumed can be controlled. Many sick neonates do not
receive colostrum shortly after birth and so are deprived of
its growth-promoting factors. The colostrum-deprived piglet
model permits the formula to be tested on stressed animals.
The restrictions and limitations of research projects with
foals and infants encourages the use of an appropriate
animal model for all preliminary work.

48
This study was part of a multi-project program to
develop an enteral diet for premature or sick foals.
Designing such a diet is a complex task requiring a series
of separate trials to determine the ideal formulation.
The goals of this specific study were:
1. to learn techniques and methods used in the investigation
of neonatal nutrition using a piglet model;
2. to determine the effect of an elemental, enteral diet on
the growth, intestinal development and general health of
colostrum-deprived neonatal piglets and, secondarily, to
learn to formulate diets with different lipid sources;
3. to evaluate and improve surgical and daily care
procedures for colostrum-deprived piglets hand-raised for 7
d; and,
4. to use the information obtained from the piglet study
towards possible development of a diet for neonatal foals.
Materials and Methods
Animals
This study was conducted using newborn Yorkshire x
Hampshire x Duroc crossbred piglets. The sows were bred and
cared for by the staff of the Reproductive Physiology Unit
under the direction of Dr. Fuller Bazer of the Animal
Science Department. Labor was induced 2 d prior to the
expected farrowing date to ensure that the birth would be
attended. The sow was injected with Prostaglandin F2a at

49
730 h on d 111 of gestation. Oxytocin was administered at
900 h on d 112. Farrowing usually occurred within several
hours after the oxytocin injection. All piglets were
removed from the sow immediately after birth, before
nursing. The umbilical cord was tied and cut. Identifying
markings, sex, weight, and general condition were recorded.
Only piglets greater than 750 g were admitted to the trial.
The protocol that subseguently was followed depended on the
treatment assignment of the litter.
Piglets from a total of 10 different litters were used
for the trial. Only one litter was on trial at any one time
due to time, labor, and facility constraints.
Baseline values were obtained from 18 piglets from two
litters removed to the animal facilities at the Food Science
and Human Nutrition Building and euthanized within a few
hours after birth. These non-fed piglets are referred to as
birth piglets (B0).
Normal values for sow-fed animals (SF) were obtained
from 22 piglets from two litters were returned to the sow
and allowed to nurse for 7 d, after which they were
euthanized. The sow and piglets were housed in an
environmentally controlled room. The sow was confined to a
farrowing stall of pipe construction. The piglets could
enter the stall easily under the bottom pipe or retreat out
of the stall to a corner of the room to avoid the sow.
Rubber floor mats were used to reduce conductive heat loss

50
and improve footing. Clean hay was provided for bedding in
a corner of the room where the piglets often slept. A heat
lamp provided additional warmth. The SF were monitored and
weighed daily.
Thirty-eight experimental piglets from six litters were
transported to the Piglet Neonatal Intensive Care Unit
(PNICU), an environmentally controlled room (37°C, 70%
humidity) similar to hospital nurseries designed for human
neonates. These colostrum-deprived piglets were hand-raised
in individual plexiglass boxes (measuring 14" x 14" x 17")
in the PNICU for 7 d.
Surgery
All experimental piglets except those bottle-fed the
milk-based diet underwent surgery. Umbilical artery and
bladder catheters were inserted to permit collection of
blood and urine, respectively, during the trial. The
samples were used to evaluate the health of the piglets.
Gastric catheters were inserted to permit feeding by
constant infusion pump.
Development of anesthesia procedures. Piglets were
anesthetized for surgery with intramuscular injection (75
mg/kg BW) of ketamine HC1. Lidocaine was given
subcutaneously at the incision site. Once an intravenous
(i.v.) catheter was placed in the umbilical artery, an
Acepromazine drip was started at 60 drops/min. Full

51
recovery from this anesthesia was slow, delaying the
initiation of enteral feeding.
In an effort to decrease recovery time, the last two
litters on the trial were given the inhalant isoflurane by
mask as the sole anesthetic. Schieber et al. (1986) found
that although isoflurane caused a reduction in blood
pressure, peripheral resistance was reduced, preventing a
decrease in cardiac output. These workers suggest that
isoflurane, well tolerated by newborn piglets, has a clear
advantage over the use of halothane. The piglets in the
present study tolerated isoflurane well, recovering quickly
(some were quite active within minutes of removal of the
inhalant).
Catheter placement. Each piglet was fitted with an
umbilical artery catheter for administration of i.v. fluids
(.45% saline and 5% dextrose) during and after surgery, and
for collection of blood on d 3 and d 5 of the trial. The
piglets received the i.v. fluids until they were fully
recovered from the anesthetic and enteral feeding was
initiated, at least 12 h post-surgically.
Piglets also were fitted with bladder catheters for
urine collection. The ureters were left open to permit
normal urination.
The introduction of gastric catheters and the use of
continuous infusion pumps allowed enteral feeding to be

52
All catheters were externalized through a single
abdominal incision.
Post-operative care. After surgery, the piglets were
returned to their plexiglass boxes in the PNICU and observed
closely. A chart was kept beside each box for recording
data from individual piglets. Respiration, temperature,
color, activity level, and urine output, were recorded every
30 minutes until the piglets had recovered fully.
Diets
Hand-raised piglets were fed one of three enteral
formulations. A conventional, cow's milk-based formula was
bottle-fed to a total of 12 piglets from two litters. These
piglets, designated Group C, did not undergo surgery. In
addition, four piglets were fed this formula via gastric
catheter. Previous trials had shown that the growth rate of
piglets fed this formula was similar to the rate of sow-fed
piglets.
The two other diets used in this study were based on
the liquid elemental formula Peptamen (Carnation). All
piglets fed diets containing the elemetal formula were
designated Group E. Hydrolyzed whey proteins provide amino
acids and malto-dextrin is the carbohydrate source. The
formula was fed in two different, isocaloric forms based on
the predominant fat source. These were designated as MCT
(medium chain triglyceride) or LCT (long chain triglyceride)

53
the predominant fat source. These were designated as MCT
(medium chain triglyceride) or LCT (long chain triglyceride)
formulas. Because Peptamen is much higher in carbohydrate
than sow's milk, it was diluted to one-half concentration
before use. However, this necessitated the addition of whey
(27.4 g/liter for the MCT formula and 35 g/liter for the LCT
formula), calcium citrate (4.38 g/liter), and calcium
phosphate (5.77 g/liter) to obtain a composition similar to
sow's milk. The wrong type of whey mistakenly was added to
the formulas fed to one of the litters, invalidating the
data obtained from these pigs. The diets resembled sow's
milk in the amount of fat (60 g), protein (75 g), and
carbohydrate (63.5 g) provided per liter. Carnitine was
added at 486 ¿il/liter of formula. The MCT formula was
obtained by adding 15 g medium chain triglyceride oil (MCT
Oil, Mead Johnson and Co., Evansville) and 6 g sunflower oil
(SFO) to 1 liter of Peptamen. All of the lipids in SFO are
long chain fats. The final MCT:LCT ratio was 70:30. The
LCT formula required the addition of .8 g MCT oil and 51.8 g
SFO to 1 liter of low-fat (LF) Peptamen. This resulted in a
90:10 LCT:MCT ratio. The composition of sow's milk and the
dietary treatments used in this trial are presented in
Tables 2.1-2.2.

Table 2.1 Composition of dietary treatments, base formulas, and sow's milk
on a per liter basis.
Diet
Composition
Protein
Fat
Carbohydrate
Calcium
Phosphorus
g
kcal
g
kcal
g kcal
g
g
Sow's milk
72
288
61
549
48 192
2.1
1.9
Peptamen
(U-5122-1)a
40
160
39
351
127 508
VO
•
o
0.5
Low Fat
Peptamen
(U-5216)b
47.6
190
7.4
67
186 744
0.6
0.5
MCT Formula
75
300
60
540
63.5 254.5
2.5
2.0
LCT Formula
75
300
60
540
93 372
2.5
2.0
Base formula
used
for MCT
diet.
DBase formula
used
for LCT
diet.

55
Table 2.2 Composition of sow's milk (Pond and Houpt, 1978)
and Peptamen (Carnation)
on a per liter basis.
Nutrient
Sow's Milk
Peptamen
Protein, g
72
40
Fat, g
61
39
Carbohydrate, g
48
127
Folic Acid, g
3900
4000
Thiamin, g
650
1500
Riboflavin, g
1370 -
1700
8200
Niacin, mg
4.3 - 9.0
20,000
B6 • 9
200
3000
-6
B12, g
1.4
6 X 10 °
Biotin, g
14
.0003
Pantothenic
Acid, mg
4
10
Vitamin A, IU
500 - 8500
3750
Vitamin D, IU
100
200
Vitamin E, IU
1.4
20
Vitamin C, mg
146
100
Calcium, mg
2100
600
Phosphorus, mg
1000 -
500
1900
Potassium, mg
1000
1250
Sodium, mg
340
500
Magnesium, mg
200
300
Chloride, mg
1000
1000
Iron, mg
1.33
9
Zinc, mg
4.94
10
Manganese, mg
2.0
Copper, mg
1
Iodine, mg
.075
Energy, kcal
1030
1000

56
Development, of Feeding Regimen
The elemental diets were fed by bottle to the first
litter of piglets that received these formulas. Subsequent
piglets fed these diets were fed via gastric catheters. The
formulas were fed to the bottle-fed piglets at the rate of
15 ml/h on d 1 and 35 ml every 2 h on d 2. The rate was to
be increased by 5 ml/feeding on each successive day,
remaining at 50 ml/2h on d 6 and 7. Because the piglets in
this litter suffered from severe, watery diarrhea, the
formula was changed to Peptamen alone, at two-thirds
strength on d 5. Subsequent litters of Group E piglets, fed
via gastric infusion, received 10 ml ' h-1 ' kg BW-1 on d
1. On each successive day, the amount fed to healthy
piglets was increased by 3 ' h-1 ' kg BW-1. The flow rate
was not increased for piglets that were not thriving. The
last two litters were fed the MCT and LCT diets at half¬
strength. This decreased the production of watery feces.
With ketamine anesthesia, long post-surgery recovery
times delayed oral feeding. These piglets were often slow
to show normal sucking and swallowing behavior after being
anesthetized. Placement of a gastric catheter during
surgery allowed commencement of enteral feeding much sooner
than was possible with bottle-feeding. Enteral nutrition
was initiated on an individual basis when the piglet
appeared to be awake and kicking motions of the hindlegs
were observed. Usually, these animals remained on i.v.

57
fluids post-surgically for about 8 to 12 h before receiving
enteral nutrition.
To determine the effect of PNICU conditions on piglets,
the Group C controls were bottle-fed a milk-based formula
designed to be similar to the composition of sow milk. To
determine the effect of surgery on PNICU piglets, four
animals (surgery controls) were fed the milk-based diet via
gastric catheters using a continuous infusion pump. The use
of infusion pumps reduced the variation in post-absorptive
state that might occur when the animals are euthanized at
different times.
In summary, a total of 13 Group E piglets were fed the
elemental diets. Four piglets were bottle-fed, while the
rest received enteral nutrition via gastric catheter. Two
piglets were fed the base formula, Peptamen, with no
additions, via gastric catheter, to determine if the
additions of calcium and phosphorus were contributing to the
diarrhea. Of the 16 piglets fed the milk-based control
formula, 12 were bottle-fed and four underwent surgery for
gastric catheter placement and were fed via constant
infusion pump.
Daily Care
The piglets were monitored continuously throughout the
trial. Urine was collected asceptically from the bladder
catheters every 30 min. At morning and evening "rounds" the

58
incision site was checked and swabbed with an iodine
solution, the venous catheter was flushed, bandages changed,
and temperature, respiration, and girth measurements were
recorded. When piglets were fed via gastric catheter, new
bags of formula were started after rounds. Piglets were
weighed during morning rounds. On d 3 and d 5 blood was
collected from the umbilical catheter.
Every effort was made to keep stress on the piglets to
a minimum. Each piglet was provided with a rubber baby-
bottle nipple taped to the inside of the box, a plastic
ball, and a soft cloth.
Tissue Collection
At sacrifice, the animals were anesthetized with Na
pentobarbital and then exsanguinated by cardiac puncture.
Organs were removed as quickly as possible, weighed, and
frozen for later analyses. The intestines were measured,
weighed, cleaned of contents, and then re-weighed. The
small intestine of each piglet was divided into three
segments of equal length, designated proximal (top)
intestine (TI), middle intestine (MI) and distal (bottom)
intestine (BI).

59
Analyses
Leukocyte (WBC) counts were performed using a Unopette
dilution chamber and a hemocytometer. Microhematocrit (Hct)
determinations performed shortly after blood was collected.
Hemoglobin (Hb) was determined using the modified
Drabkins method with a Beckman DU-7 spectrophotometer at a
wavelength of 540 nm.
The plasma glucose was assayed using Trinder reagent
(Sigma Chemical Co., St. Louis), based on the glucose
oxidase reaction. The OD was read with the Beckman DU-7
spectrophotometer at 505 nm.
Non-collagen protein in intestinal homogenates was
determined using the micro-Biuret method (Itzhaki and Gill,
1964) after precipitation with .2 N NaOH. The OD was
measured with a Gilford spectrophotometer at 310 nm.
The activities of the intestinal disaccharidases
maltase, lactase, and sucrase were determined using a
slightly modified technique by Dahlqvist (1964).
Preparation of the tissue included homogenization in a
glass-on-glass hand homogenizer with .05 M sodium phosphate
buffer in a 1:5 weight/volume ratio. Each tissue homogenate
was incubated with each substrate to determine the
micromoles of glucose produced by the disaccharidase enzyme
per microgram tissue, during a 1 h incubation. The OD due to
the glucose produced was read at 505 nm with the Beckman DU-
7 spectrophotometer.

60
Urine was collected from Group E piglets every 30 min
throughout the trial. All of the urine collected during a 6
h period was pooled, unless it was unusual in appearance.
In this case, the abnormal urine was placed in a separate
vial. The urine was kept on ice or refrigerated until
analysis.
The volume of urine collected as well as a description
of its general appearance were recorded for each piglet for
each 6 h period. Fresh urine was analyzed using Ames
Multistix SG reagent strips (Miles, Inc., Ames). These
strips provided a quick assessment of specific gravity, pH,
protein, glucose, ketones, bilirubin, blood, and urobiligen.
The remaining urine was frozen, pending further analysis.
The following equation was used to calculate the % BW
gained by the 7-d old piglets:
% BW wt gain = ((d 7 BW - birth BW) -s- birth BW) * 100.
The percent of body weight of each section of the small
intestine (TI, MI, and BI) was calculated using the
following equation: SI section wt(% of d 7 BW) =
(SI section wt t d 7 BW) * 100. These results will be
referred to as %TIWT, %MIWT, and %BIWT.
Additional Information
Additional information and details of surgery, daily
care, necropsy procedures, and tissue analyses are described
by Baltzell (1988). The PhD dissertation involved the the

61
colostrum-deprived neonatal piglet model that was further
developed in this current work.
Statisical Analysis
Treatment means for SF and BO data only were obtained
by method of least squares ANOVA (Snedecor and Cochran,
1969) using the GLM procedures of the SAS Statistical
Software (Freund and Littell, 1981). The model included
treatment, litter(treatment), sex, sex*treatment, and
sex*litter(trt). Data from Group C and Group E treatments
were not analyzed because of the low numbers of animals in
each group.
Results
Numerical results are expressed as least squares means
± SE, unless stated to be observed means.
General Observations
Sow-fed piglets. All of the SF piglets were active,
healthy and growing well at the end of the 7 d trial period.
They did not seem unduly stressed by the daily weighing
procedure.
PNICU piglets. Thirty-eight piglets were started on
trial. All of the Group C piglets werew healthy and active
through by d 7. Although the elemental diets were
formulated to be similar to sow milk, the Group E piglets
did not thrive. These piglets experienced diarrhea that may

62
were made for each litter. These changes reduced but did
not eliminate the diarrhea. The survival rate of Group E
piglets was low, as only nine of 20 survived for 7 days.
This was probably due, in part to diet and to dislodged or
faulty catheters. Piglets that had dislodged gastric or
bladder catheters were humanely euthanized. Because of the
low survival rate of the Group E piglets, only descriptive
results will be presented.
Occasionally, piglets were born with umbilical hernias.
This was not generally not a problem, except for one piglet,
who diet after a portion of the small intestine
strangulated. Blockages or strictures of the small
intestine were observed in four animals.
Behavior of PNICU piglets. The hours of care required
by the piglets allowed for close observation of their
behavior. Each piglet had a particular activity pattern
that was known to all the caretakers. Therefore, early
detection of problems was possible.
The non-surgery piglets, bottle-fed the milk based
diet, quickly became accustomed to the routine in the PNICU.
They were noisy and wanted attention from the caretakers.
Even during the first day, they began to recognize the pre¬
feeding routine. They had strong suckling reflexes. Play
was part of their daily activity, just as it was for the
sow-raised piglets. Although they were kept in individual
boxes, the plexiglass allowed them to see each other and the

63
boxes, the plexiglass allowed them to see each other and the
caretakers. Objects such as crumpled paper towels, plastic
balls, and pacifiers were used as toys.
The healthy surgery piglets also exhibited play and
attention-seeking behavior and did not seem hampered by the
i.v. or enteral infusion lines attached to them. Soft
cloths were used as bandages to cover the incision and keep
lines in place. These were ignored by the animals. Swivels
in the infusion lines kept the twisting and tangling to a
minimum. These lines were changed twice daily, when fresh
formula bags were set up.
Activity level (sleeping, lying down, alert, moving,
etc.) of each piglet was recorded on the individual charts
every half hour and changes from the previous record noted.
Nasal oxygen was available, if needed. An attempt was made
to cool (water or alcohol swabs) or warm (heating pad or
heat lamp) animals whose body temperatures were not in the
normal range.
Healthy animals generally produced an adequate amount
of relatively clear urine. Animals that had decreased urine
output or bloody urine generally had enlarged kidneys on
necropsy and often had fluid accumulation in the abdomen and
hindlegs.
In an effort to improve the survival and growth rate of
the piglets fed the elemental diets, modifications in the
formulas and daily intakes were made for each successive

64
litter. After the first litter, the calcium and phosphorus
additions were discontinued. The amount of whey added to
the base formulas was decreased. The formula strength and
flow rate during the first few days after surgery were
decreased. It was thought that these changes might decrease
the diarrhea, improve tolerance of the formula, and smooth
the transition to enteral nutrition after surgery.
Gross Observations at Necropsy
Sow-raised and BO piglets. There were no abnormalities
noted in the organs of any of the SF or the BO piglets. The
intestinal tissue of the BO piglets was very thin compared
with that of the SF and could be easily torn when separating
the mesentery from the small intestine.
The stomachs of the BO piglets essentially were empty,
while the stomachs of SF piglets contained milk curd. The
proximal intestinal contents of the SF piglets were very
liquid and yellowish. The contents became thicker in the
more distal areas. The cecal contents were pasty and darker
yellow. Large intestine contents were very thick and became
darker in the more distal regions. The distal colon
contained formed fecal pellets, dark in color.
Post-surgical, formula-fed piglets. In all cases, the
positions of the catheters were verified on necropsy. No
abnormalities, other than occasional adhesions, were noted

65
with the catheters. The lining of the stomachs and bladders
did not show irritation from the catheters.
Some piglets exhibited respiratory distress and had
reddish, congested lungs at necropsy. Some of these animals
had displayed swollen shoulders, hindlegs, and abdomens with
bluish blotches seen beneath the skin. While cause of these
blotches is unknown, it may have been due to caretakers
holding the piglets too tightly during rounds.
Many animals with bladder catheters had enlarged
kidneys. In some cases, the ureters also were enlarged.
The livers of all of the piglets appeared normal, but
several piglets had gall bladders that contained a dark,
thick material. This was not seen in the SF or BO groups.
Non-surqical, formula-fed piglets. The Group C
piglets, fed the conventional, milk-based diet, were healthy
and active at the end of the 7 d. The organs of these
piglets appeared normal at necropsy.
Some piglets bottle-fed the elemental formulas had
slightly reddish, congested lungs.
Weight Gain
The means reported in this section are observed means ±
SE.
Sow-fed litters. The sow-fed piglets (SF) increased
their birth weight by 65.75%. One litter (n=13) gained an
average of 74.49% ± 4.24, with a minimum of 50.76% and a

66
maximum of 115.18%. The other litter (n=9) gained an
average of 53.13% ± 5.90. The least amount gained was
27.58%, and the most 79.16%.
Formula-fed piglets. The birth weight of Group C
piglets (n=12) increased by an average of 78.56% during the
week long trial. The smallest gain was 54.40%, while the
largest was 96.41%. The surviving surgery controls, fed the
milk-based diet via gastric catheters (n=2), gained only
29.75%.
One piglet fed one of the MCT-based diets gained 5.63%
of its birth weight and one piglet increased its weight by
21.91%. The other piglets experienced losses of 6.89%,
11.09%, and 8.5%. The MCT-fed piglets were all very thin by
the end of 7 d. None of the four surviving piglets fed
one of LCT-based lost weight, but they gained very little.
The average gain was 11.99%. The two bottle-fed piglets
gained 6.65% and 1.85% of their birth weight. The other
two, fed by gastric infusion, gained 35.29% and 4.18%. All
but two Group E piglets weighed greater than 1 kg at birth,
and these exceptions were over 900 g.
Small Intestine Length
The total length of the small intestine (SI) was
measured at necropsy. The length of the SI in the B0
piglets was used as a reference baseline value.

67
The least squares means (LS means) for one litter of BO
and one litter of SF piglets were 284.39 cm ± 11.27 and
371.73 cm ± 9.34, respectively.
Blood Glucose
Blood glucose may be expected to vary depending on an
animal's nutritional status, health, and physical condition.
The use of infusion pumps may have reduced variations due to
differences in times after feeding at death. This was
difficult to control with bottle feedings due to time and
labor constraints. The results in this section are reported
as observed means unless stated otherwise.
The LS means of blood glucose concentration of two
litters of SF piglets and one litter of BO piglets were
13.37 mM ± 1.81 and 6.55 mM ± 2.03, respectively.
Small Intestine Weight
The %TIWT, %MIWT, %BIWT observed for BO piglets, and
sow-fed piglets are presented in Table 2.3. The BO mean was
obtained from data from one litter and the SF mean was
obtained from two litters.
Disaccharidases in the Small Intestine
Results of the disaccharidase analyses are calculated
based on the amount of glucose produced by 1 g of tissue as
a result of substrate hydrolysis occurring during a 1 h

68
incubation period. The means for the enzyme activities for
SF and BO piglets are presented in Tables 2.4-2.6. The
results for the SF and BO piglets are from one litter of
each group. Therefore, the standard errors in the tables
also include litter effects.
Group C piglets (n=12), fed a milk-based formula, and
the Group E piglets had very low intestinal sucrase
activities. In tissue samples from several Group E piglets
no sucrase activity was detected.
Maltase activity was detected in Group E piglet
intestines, as well as in tissue obtained from Group C
piglets even though the Group C piglets had not ingested
maltose. Similarly, lactase activity was detected in the
Group E piglets that had never ingested lactose.

69
Table 2.3. Least squares means for the proximal, middle, and
distal sections of the SI, expressed as a
proportion of body weight.
Treatment
%TIWT
SE
%MIWT
SE
%BIWT
SE
Sow-fed (SF)
.83
.03
.84
. 03
.87
. 03
Birth (BO)
CO
in
•
. 04
. 63
. 03
. 62
. 04

Table 2.4. Least squares means of sucrase activity in proximal, middle, and distal
sections of the small intestine of 7-day old sow-fed piglets and
piglets killed at birth.
Sucrase Activity
nmol glucose produced/h/g wet tissue
TI
MI
BI
Treatment
Mean
SE
Mean SE
Mean
SE
BO
1.2
. 85
<.l .94
1.2
.70
SF
6.6
1.18
3.2 1.31
. 6
.98

Table 2.5. Least squares means of maltase activity in proximal, middle, and distal
sections of the small intestine of 7-day old sow-fed piglets and
piglets killed at birth.
Maltase Activity
/Ltmol glucose produced/h/g wet tissue
TI
MI
BI
Treatment
Mean
SE
Mean
SE
Mean
SE
BO
14.1
3.84
14.2
3.74
11.7
.93
SF
44.2
4.22
46.7
5.19
23.2
1.29

Table 2.6. Least squares means of lactase activity in proximal, middle, and distal
sections of the small intestine of 7-day old sow-fed piglets and
piglets killed at birth.
Lactase Activity
Mmol glucose produced/hour/gram wet tissue
TI
MI
BI
Treatment
Mean
SE
Mean
SE
Mean
SE
BO
45.1
8.89
217.2
24.23
82.3
8.07
SF
34.4
12.35
171.3
33.65
74.4
11.21

73
Complete Blood Counts
The complete blood counts (CBC), consisting of Hb, Hct,
and white blood cell counts, were performed to determine if
the blood components measured fell within published normal
ranges for piglets of this age. The CBCs provided a method
of monitoring the general health of the piglet during and
after 7 d on the trial. However, it was difficult to
collect blood from the umbilical artery catheter on d 3 and
5 of the trial as the catheters often became blocked and
only a very small amount of blood, if any, could be
withdrawn. Not enough data was collected on these days to
provide meaningful results. Data from SF and BO piglets is
presented in Table 2.7.

74
Table 2.7. Results of CBCs from blood collected at
necropsy via cardiac puncture from sow-fed
piglets and piglets killed at birth.
Treatment
na
Hb
(g/di)
Hct
(%)
WBC
(cells/ml3)
SF
22
5.6-10.5
18.7-35.3
7000-18,700
BO
9
3.3-20.5
10.3-42.2
4050-10,100
7_d h
sow-fed
31
3.6-5.2
24.1-34.9
8900-12,700°
i-d b
sow-fed
70
12-12.7
39.6-43.5
6270-17,600
anumber of
samples
obtained
u
DSchmidt and Tumbleson, 1986
cd 28 data

75
Urinalysis
After surgery to introduce a bladder catheter, the
piglets routinely had some amount of blood in the urine.
Piglets with low urine outputs tended to have urine than was
visibly bloody, while piglets with substantial urine output
tended to have clear, yellow urine that was found to contain
blood only by chemstrip analysis.
The urine routinely contained varying amounts of
protein. This also may be expected after surgery. The pH
of the urine varied among piglets, but an individual piglet
tended to have urine in a small pH range throughout the
trial. The apparent health of the piglets did not seem
related to the changes in urine pH or specific gravity.
Discussion
The purpose of the work using piglets was to learn to
develop techniques and procedures to be used in a trial
designed to evaluate an elemental, enteral diet for sick or
premature neonates. To achieve this, two elemental formulas
and one milk-based formula were fed to the neonatal
colostrum-deprived piglets for 7 d. The elemental formulas
differed in the amount of fat provided by LCT and MCT
lipids. The elemental diets contained malto-dextrin as the
carbohydrate source, rather than lactose because compromised
neonates may have reduced lactase activity.

76
The animals in the first litter were bottle-fed the
elemental diets. None of them appeared to thrive during the
7 d study period, probably due to diarrhea and respiratory
problems. All subsequent Group E piglets were fed via
gastric catheters.
The diets were modified for each litter in an attempt
to develop a formulation suitable for the piglets. Changes
in the diet and flow rate reduced but did not eliminate the
diarrhea. Some workers had suggested that maltase may be
low in neonatal piglets. Therefore, it was likely that
undigested maltose reaching the large intestine may have
caused the diarrhea. This led to the investigation of the
carbohydrate source, maltose.
Disaccharidases in Neonatal Piglets
Human infants have adequate maltase at birth to digest
maltose even when they are born prematurely (Paige and
Bayless, 1981). However, there are conflicting reports in
the literature as to when maltase becomes active in the
newborn piglet. Very low maltase activity activities have
been detected in the small intestine of piglets as early as
2 d of age (Dahqvist, 1961a), and the activity appears to
increase from d 3 to 10 (James et al., 1987). The maltose
fed to the Group E piglets was within the amount reported to
be digested by normal, sow-fed piglets, but it is possible

77
that these colostrum-deprived piglets lacked sufficient
total tract maltase activity to meet their energy needs on a
maltose-containing diet.
Investigation of the tissue disaccharidase activities
in the small intestine of the experimental piglets was
undertaken to determine the ability of newborn and 7-d old
piglets to digest sucrose, maltose, and lactose.
Lactase activity was measured for reference as this
enzyme should be very active during the first week after
birth. Sucrase also would be a useful reference because
only low activities have been reported in 1-wk old piglets
(Veum and Mateo, 1986) .
The large variation in disaccharidase activities
between piglets in this trial was also reported by othe
workers (Manners and Stevens, 1972). It is difficult to
interpret the data when the large variation is coupled with
low numbers of animals per treatment.
The lactase activity along the SI was similar for the
BO and SF piglets. This is to be expected, because mammals
must be prepared, at birth, to digest lactose. The Group E
piglets, although not thriving, did have lactase activity,
even though they never consumed lactose. The intestine
remained prepared for the ingestion of this sugar throughout
the week.
The BO piglets had low maltase activities, leading to
the conclusion that piglets may not be able to digest

78
maltose efficiently enough to meet their energy needs when
this sugar is the sole carbohydrate source. The sow-fed
piglets appeared to have a numerically higher maltase
activity than did the piglets killed at birth even though
the SF piglets had never ingested maltose. This suggested
that age is a factor in enzyme development. Birth and
feeding, especially of colostrum, have been shown to
stimulate enzyme and intestinal development (Widdowson,
1985).
Sucrase activities were very low, compared with the
other disaccharidases measured. Dahlqvist (1961a) found no
appreciable sucrase in 2-d old piglets, but Beilis (1957)
reported equal digestion of lactose and sucrose by d 3.
Manners and Stevens (1972) were barely able to detect
sucrase activity until the piglets were 1 wk old. James et
al. (1987) found that EGF increased sucrose in neonatal
piglets. EGF is secreted in colostrum. In the present
study, only the SF ingested colostrum and this may explain
the generally higher enzyme activities measured in this
group.
Growth Rate in Neonatal Piglets
The Group C controls grew at a rate comparable with,
that of the sow-fed piglets. While the hand-raised piglets
did not have to expend energy to maintain body temperature
or compete for food, they did not have the benefit of

79
colostrum or sow's milk. Even though the growth rate of
Group E piglets was poor, the growth rate of the Group C
piglets indicated that colostrum-deprived piglets can be
raised successfully raised for 7 d in an intensive care
facility. The unit used in this study was kept extremely
clean but not sterile.
Development of the Small Intestine in Neonatal Piglets
It was suggested that the ingestion of colostrum and
milk affect the growth and development of the small
intestine in newborn mammals (Widdowson, 1985). An ideal,
enteral formula should supply adequate nutrients and growth
factors, especially to the small intestinal tissue. The
piglets killed at birth had very small, fragile intestines
that could be distinguished easily from the intestines of
older, fed piglets. By 1 wk of age, the small intestines of
SF piglets were longer and heavier than the BO intestines.
Because these weights are on a wet tissue basis, it is not
known whether the heavier weights are due to fluid or dry
matter.
Blood glucose
Although blood glucose is regulated by insulin and
glucagon, it can be affected by nutritional state, general
health, and time after feeding. Piglets are born with low
energy stores that must be replenished shortly after birth.

80
Blood glucose was low in BO piglets. These piglets often
were not sacrificed for several hours after birth, so
partial depletion of their energy stores probably occurred.
Normal blood glucose for neonatal piglets has been
reported by Tumbleson and Schmidt (198 6) . The values had
large standard errors, which is expected as glucose varies
with time after feeding. They reported mean values for
birth and 1-wk old piglets of 2.67 ± 1.06 and 6.67 ± 1.11
mM/L, respectively. The birth value is lower than the 6.63
± 1.54 mM measured in piglets killed shortly after birth in
this study. All of the Group E piglets had blood glucose
concentrations similar to the reported values for 1-wk old
piglets. However, the 1-wk old SF and Group C piglets had
concentrations averaging about 13 mM. Method of blood
collection, stress, nutrition, breed effects, and time of
sampling have been known to affect results (Tumbleson and
Schmidt, 1986) .
Hematology
Results of the CBCs were compared with values reported
by Schmidt (1986). Hemoglobin in neonatal piglets was
reported to be 12.5 ± .2 g/dl at birth, and 10.0 ± 1.8 g/dl
by d 7. The Hct values at birth are 40.3 ± .7% and 29.5 ±
.4% by d 7. The piglets in the present study had
considerable variation in the Hb and Hct values, but most
were within or close to the reported ranges. However, the

81
PNICU surgery controls (n=2) had low Hb and Hct values.
These piglets had been very weak and may have had problems
unrelated to the treatment or surgery.
At birth, piglets would be expected to have about 6000-
1700 WBC//U1, and 7000-10,000 WBC//il by d 28 (Schmidt, 1986).
Generally, the piglets that survived until d 7 had CBC
values that were close to the normals for their age. Many
of the Group E piglets had severe diarrhea but did not have
extremely high packed cell volumes, suggesting that the
formula flow rate was sufficient to prevent dehydration.
Summary of Piglet Trial
As a result of this study, some techniques and
procedures for performing trials on colostrum-deprived
piglets were refined. Although each successive study will
probably result in further improvements, the colostrum-
deprived piglet model currently can be used successfully to
investigate the nutritional needs of neonates.
The poor growth rates of the piglets fed the MCT and
LCT diets may have been due to the carbohydrate source. No
information regarding the benefits of the addition of MCT to
neonatal formulas could be obtained from this trial.
However, using an appropriate base formula should allow this
information to be obtained from future studies using the
piglet model.

82
deprived piglet model currently can be used successfully to
investigate the nutritional needs of neonates.
The poor growth rates of the piglets fed the MCT and
LCT diets may have been due to the carbohydrate source. No
information regarding the benefits of the addition of MCT to
neonatal formulas could be obtained from this trial.
However, using an appropriate base formula should allow this
information to be obtained from future studies using the
piglet model.
It is clear that only digestible carbohydrates should
be included in neonatal enteral formulas. However,
information on the digestibilities of various sugars may not
be available for each species. Development of an enteral
diet for foals based on piglet data was a goal of this
research. However, it was not known whether newborn foals
had sufficient maltase activity to adequately digest the
malto-dextrins in the base elemental diet. Therefore, an
oral disaccharide tolerance test was performed on neonatal
foals.

CHAPTER III
ORAL DISACCHARIDE TOLERANCE TEST
FOR NEONATAL FOALS
Introduction
Neonates that are sick, stressed, or born prematurely
may have poorly functioning digestive systems. The pattern
of small intestine disaccharidase development may not be the
same for these newborns as for healthy neonates (Rossi et
al., 1986). Lactase activity can be reduced greatly as a
result of prematurity or disease (Rossi et al., 1986;
Tzipori et al., 1984). Consequently, lactose may not be
suitable for use in enteral diets designed to support non-
healthy neonates. The use of other carbohydrate sources
needs to be considered carefully. Including monosaccharides
in enteral diets would increase the osmolarity of the
formula higher than neonatal intestines could handle well,
possibly causing diarrhea. Complex sugars must be cleaved
into smaller and smaller units, and finally from
disaccharides into single sugar molecules before absorption
can take place. It is not known if neonatal foals can
digest and absorb disaccharide sugars other than lactose.
Therefore, it is important to determine which enzymes are
likely to be active in the intestine of equine neonates.
83

84
Enzyme activity varies with age and species and can vary
among similar individuals (Paige and Bayless, 1981). There
is very little information in the literature about the
disaccharidase activity in the small intestine of foals,
although it was studied in neonatal humans, pigs, calves,
and rats (Rossi et al., 1986; Dollar and Porter, 1957; Code,
1968; Paige and Bayless, 1981).
Dollar and Porter (1957) concluded from their study of
newborn calves that enzyme activities determined from
mucosal cell extracts were correlated closely to the results
of an oral disaccharide tolerance test. Using a similar
procedure, Roberts et al. (1974) studied lactase, maltase,
and sucrase activities in horses of various ages by
homoginizing samples of intestinal mucosa. Small intestine
tissue was obtained from the horses slaughtered or
euthanized. However, the data does not show the pattern of
disaccharidase development during the critical first week
postpartum. Lactase was detected as early as 105 d
gestation. Peak activity was found in foals at birth and it
remained high for 4 mo. Sucrase was barely detected in
fetuses from 3 to 9 mo gestation, remaining low at birth.
Adult activities were attained by 7 mo of age. Similarly,
maltase was very low until the ninth month of gestation.
Activities at birth were 12 to 15% of the enzyme activities
of adult equines. The authors reported a gradual increase,
until adult activities were reached at 7 mo of age.

85
In a subsequent study, Roberts (1975) determined the
ability of horses to digest and absorb disaccharides using
an oral disaccharide tolerance test. These results agreed
with those of his earlier study. However, this study did
not include data on very early neonatal life.
A non-invasive method of estimating disaccharidase
activity, such as an oral tolerance test, would allow foals
to be studied at more than one point in time and would not
require the experimental animals to be euthanized.
The present study was done using healthy foals, nursing
their mothers. Although the results will not necessarily be
applicable to sick or premature foals, it may eliminate some
disaccharides as possible ingredients for enteral diets; if
healthy foals are unable to digest a particular
disaccharide, then it is unlikely that a stressed foal would
have that ability. The results of this study will provide
information on the digestive ability of newborn, healthy
foals non-invasively, and help determine possible sources of
carbohydrates for use in enteral formulas. Because the
animals do not need to be euthanized to obtain samples, each
foal can be used on different days to determine the changes
in enzyme activity during the first week of life. There can
be considerable variation in activities among animals
(Manners and Stevens, 1972) .
Selection of a carbohydrate source for inclusion in
elemental, enteral formulas will depend on the activity of

86
the necessary enzymes and the desired osmolarity of the
formula. Disaccharidases are important in the digestion of
all sugars larger than monosaccharides.
It was presumed that healthy foals would have adequate
lactase activity while relying on milk for nourishment. It
was also presumed that glucose would be easily absorbed by
these neonates. Therefore, the changes in plasma glucose
occurring after administration of glucose and lactose would
be used for comparison with test sugars. Maltose and
sucrose were chosen as test sugars. Maltose was included
because it was the carbohydrate source in the base formula
fed to the piglets. Sucrose was tested because it was
thought that this sugar would be poorly digested and would
provide a reference for a non-absorbed substrate.
The goals of this trial were:
1. to investigate the ability of newborn, healthy
foals to digest maltose, lactose, and sucrose during the
first 5 d postpartum;
2. to investigate the change in blood glucose in
healthy foals after a short fast and the effects of age on
the ability of the foals to maintain blood gluocse
concentration after a short fast; and,
3. to determine the changes in blood glucose in
healthy foals in response to an oral glucose challenge.

87
Materials and Methods
Preliminary Study
A preliminary study to determine the effect of nursing
on blood glucose was performed on a normal, healthy foal at
8 h postpartum. Care of the mare and foal during and after
birth followed standard procedures at the University of
Florida Horse Research Center.
At 8 h of age, the foal was fitted with a Teflon
jugular vein catheter (Quik-Cath, Travenol Laboratories,
Inc.), in an aseptic procedure, to allow periodic blood
sampling. To minimize stress to the foal, a small amount of
local anesthetic was injected at the catheter site. The
plastic wings of the catheter were fixed to the skin with a
superglue adhesive. This was easily removed with acetone at
the conclusion of each trial period. The catheter was
filled with sterile, heparinized saline between samplings.
The foal was then allowed to return to the mare and resume
normal activity. After the foal nursed the mare, the time
was noted and the foal was muzzled to prevent nursing.
Blood was drawn from the catheter at this time (time = 0).
Blood was then drawn every 30 min thereafter for 2 h. This
procedure was repeated on the same foal on d 5. This study
suggested that blood glucose reached a baseline
concentration after a 1 h fast. It was concluded that a 2 h
fast prior to administration of the treatment, and a 4 h
fasting collection period would be sufficient to monitor

88
changes in blood glucose after a test meal in the subsequent
experiment.
Animals and Design
Eleven Quarter Horse and two Thoroughbred foals were
used for the main trial. The foals remained with their dams
throughout the trial. Foals were weighed shortly after
birth and on d 5. After each foal was born, it was randomly
assigned, by sex, to one of four treatment groups until the
block was filled. This procedure was repeated for
subsequent foals, assigning foals of each sex to fill the
treatment blocks evenly.
On d 1, at 6 h postpartum, each foal was fitted with a
jugular vein catheter, as described above. Foals were
muzzled to prevent nursing. The foals were then fasted for
2 h to allow blood glucose concentrations to reach a
baseline level. Each animal was then given the appropriate
treatment consisting of oral administration of either
maltose, lactose, or sucrose at 1 g (5.56 mmol) per kg BW in
a 20% solution (w/v), or a glucose solution at .5g (2.78
mmol) per kg BW. These dosages were similar to dosages used
by Roberts (1975) .
The solution was offered to the foal in a bottle fitted
with a suitable nipple. If the foal did not voluntarily
consume the solution, it was administered via dosing syringe

89
or via nasogastric tube. Blood was collected from the
catheter immediately before the solution was given
(time = 0), every 15 min for the first hour, and every 30
min thereafter for the second, third, and fourth h of the
trial period. Each foal received the same treatment when
the procedure was repeated on d 3 and d 5 postpartum.
Analyses
The blood was collected in tubes containing
ethylenediaminetetraacetic acid (EDTA), plasma removed after
centrifugation at 3500 g for 20 min and frozen. Plasma
glucose was determined using the Trinder peroxidase method
(Sigma Chemical Co.). Optical density was measured at 505
nm with a Beckman DU-20 spectrophotometer.
Statistical Analyses
Statistical analyses were performed by method of least
squares ANOVA with time as a class variable (Snedecor and
Cochran, 1969,) using the GLM procedures of SAS (Freund and
Littell, 1981). The complete mathematical model is given in
Table 3.1. The appropriate error term is shown for each
independent variable.

90
Table 3.1. Complete mathematical model for ANOVA of changes
in plasma glucose.
Source df
Treatment 3
Sex 1
Treatment*Sex 3
Foal(Treatment*Sex) 5
Day 2
Treatment*Day 6
Day*Sex 2
Treatment*Sex*Day 6
Day*Foal(Treatment*Sex) 10
Time 10
Time*Treatment 30
Time*Foal(Treatment*Sex) 50
Day*Time 20
Sex*Time 10
Day*Sex*Time 20
Treatment*Sex*Time 30
Day*Treatment*Time 60
Treatment*Day*Sex*Time 60
Error term
Foal(Treatment*Sex)
Foal(Treatment*Sex)
Foal(Treatment*Sex)
Remainder
Day*Foal(Treatment*Sex)
Day*Foal(Treatment*Sex)
Day*Foal(Treatment*Sex)
Day*Foal(Treatment*Sex)
Remainder
Time*Foal(Treatment*Sex)
Time*Foal(Treatment*Sex)
Remainder
Remainder
Remainder
Remainder
Remainder
Remainder
Remainder
Remainder
101

91
Equations for the time curves were obtained using
reduced models with time as a continuous variable. The
reduced pooled model is presented in Table 3.2. The models
for comparison of day curves, treatment curves and
day*treatment curves were similar to the pooled model except
that day*timen, treatment*timen, and day*treatment*timen,
respectively, were used in place of time11. F tests for
homogeneity of the time curves were calculated based on the
techniques described by Snedecor (1956). Rejection of the
null hypothesis would indicate that the curves differed,
i.e. were not parallel.

92
Table 3.2. Pooled regression modela for plasma glucose time
curves.
Pooled Curve
Source
df
Error term
Treatment
3
Foal(Treatment*Sex)
Sex
1
Foal(Treatment*Sex)
Treatment*Sex
3
Foal(Treatment*Sex)
Foal(Treatment*Sex)
5
Remainder
Day
2
Day*Foal(Treatment*Sex)
Treatment*Day
6
Day*Foal(Treament*Sex)
Treament*Day*Sex
6
Day*Foal(Treatment*Sex)
Day*Sex
2
Day*Foal(Treatment*Sex)
Day*Foal(Treatment*Sex)
10
Remainder
Time
1
Remainder
Time2
1
Remainder
Time2
1
Remainder
Time4
1
Remainder
Time5
1
Remainder
Time6
Remainder
1
383
Remainder
aReduced model

93
Results
General Observations
In general, the foals drank the treatment solutions
readily from a bottle on d 1 of the trial. On d 3 and 5,
some foals refused the solution. Some of these foals also
proved uncooperative in swallowing aliquots of the solution
delivered by dosing syringe. In these cases, a very small
diameter nasogastric tube was passed, usually without unduly
stressing the foal.
Some foals were uncooperative during catheterization.
These foals were allowed to relax for periods up to 1 h
after catheterization before the treatment was administered.
Throughout the trial every effort was made to keep the foals
calm. Subdued lighting, minimal restraint and keeping the
mare nearby helped reduce the foal's anxiety.
None of the foals experienced any digestive
disturbances, such as colic or diarrhea, during the trial.
The foals became adjusted to the frequent blood collections,
remaining in a recumbent position or even sleeping.
High quality reagent grade substrates were used for the
treatment solutions. Analysis of the treatment solutions
showed maltose contained 1.48% free glucose, sucrose
contained .74%, and lactose less than .01%.

94
Fasting Blood Glucose
The plasma samples collected from all foals at time = 0
were used to determine fasting glucose concentrations. At
time = 0 no treatment had been given to the foals and they
had been treated alike. Therefore, data from all foals were
pooled.
Breed and breed interaction effects were found to be
non-significant (P > .05). Rogers et al. (1983) reported
higher blood glucose concentrations in young fillies than in
colts, although results of statistical analyses were not
reported. However, we found no differences due to sex in
the 13 foals studied (P = .8957).
The LS mean of blood glucose from fasted foals was 4.14
mM on d 1. The means on d 3 and d 5 were 6.08 and 6.31 mM,
respectively. The SE of the means is .27. The mean from d
1 is lower than the mean from the other 2 d (P = .0001).
Day 3 and d 5 means were not different from each other.
Glucose Absorption Time Curves
Using time as a continuous variable, equations for
treatment, day, and treatment-day curves were calculated.
Sequential addition of higher order terms showed time to the
fifth power to be significant (P < .05). However, fifth
order equations led to negative estimates near the right
hand portion of some curves. Equations of the sixth order
were then used, as these produced logical curves. Sixth

95
order equations were used to describe all of the curves.
However, sequential addition of hiqher order terms revealed
that the linear term only was sufficient to describe the
plasma glucose time curves resulting from sucrose
administration.
The equations for the day-treatment curves are
presented in Tables 3.3-3.6. Time curve equations represent
the changes in plasma glucose due to treatment
administration. Thus, changes in plasma glucose occurring
after dosing with oral glucose are represented by the
glucose (G) time curves. Similarly, lactose (L), maltose
(M) and sucrose (S) curves represent changes in plasma
glucose in foals dosed with these sugars.
Treatment Curves Pooled Over All Days
Orthogonal contrasts and tests for homogeneity of the
curves were performed on data from each treatment pooled
over all 3 d. The L curve was not different from the M and
S curves combined. The G curve was not different from the
pooled M, L, and G curve. The M curve was different from
(i.e. not parallel to) the S curve (P = .0220). The G and L
curves also were different from each other
(P = .0186).

96
Table 3.3 Equations for day*treatment curves for lactose.
DAY 1
Ya = 5.224 + .0103Tb + 2.273xl0_3T2 - 3.979xl0-5T3
+ 2.526X10-7T4 - 7.105xl0-9T5 + 7.564xlO_13T6
DAY 3
Y = 7.013 + .162T - 3.167x10-3T2 + 1.632xlO~5T3
+ 3.606x10_8T4 - 4.982xlO_10T5 + 1.020xl0-12T6
DAY 5
Y = 6.847 + .159T - 3.036xlO-3T2 + 1.805xl0”5T3
+ 2.533xl0-8T4 - 8.952x10-1IT5 + 2.195xlO-13T6
aY = plasma glucose, mM
bT = time (continuous variable)

97
Table 3.4 Equations for day*treatment curves for maltose.
DAY 1
Ya = 5.361 + .00136Tb + 1.404xl0_3T2 - 2.634xlO~5T3
+ 1.945X10-7T4 - 6.494xlO-10T5 + 8.180xl0_13T6
DAY 3
Y = 6.192 + .076T - 1.939xlO~3T2 + 1.806xl0_5T3
- 7.001xl0-8T4 + 9.03Ixl0-11T5 + 2.322xlO-13T6
DAY 5
Y = 6.031 + .124T - 3.000x10-3T2 + 2.885xl0_5T3
- 1.381x10_7T4 + 3.145X10-10T5 - 2.535xlO~13T6
Y = plasma glucose, mM
T = time (continuous variable)

98
Table 3.5 Equations for day*treatment curves for glucose.
DAY 1
Ya = 3.020 + .167Tb - 2.041xl0_3T2 - 1.381xlO_5T3
+ 3.102x10-7T4 - 1.558x10~9T5 + 2.510xl0-12T6
DAY 3
Y = 5.762 + .228T - .0114T2 + 1.981xlO_4T3
- 1.553xlO“6T4 + 5.615xlO-9T5 - 7.623xlO-12T6
DAY 5
Y = 6.260 + .133T - 5.161xlO~3T2 + 6.919xlO_5T3
- 4.406x10-7T4 + 1.356X10-9T5 - 1.622xlO~12T6
aY = plasma glucose, mM
bT = time (continuous variable)

99
Table 3.6 Equations for day*treatment curves for sucrose.
DAY 1
Ya = 3.692 - .120Tb + 3.408xl0-3T2 - 4.227xl0-5T3
+ 2.506x10-7T4 - 7.007xl0-10T5 + 7.408xl0_13T6
DAY 3
Y = 5.480 - .001T + 8.630xl0-4T2 - 2.559xl0_5T3
+ 2.581x10~7T4 - 1.085x10-9T5 + 1.628xlO_12T6
DAY 5
Y = 6.748 + .035T - 1.299xlO-3T2 + 1.721xlO~5T3
- 1.110X10~7T4 + 3.410xl0-10T5 - 3.946xlO_13T6
aY = plasma glucose, mM
T = time (continuous variable)

100
Comparison of Day-Treatment Curves
The least squares means for each treatment for each of
the days are presented in Table 3.7.
Day 1. The M curve was different from the S curve (P =
.0059), the L curve (P = .0015), and the G curve (P =
.00002). The G curve was different from the L curve (P =
.0028) (Figures 3.1-3.2).
Day 3. The M curve was not parallel to the L curve (P
= .0027), but was not different from the G curve on this day
(Figures 3.3-3.4) .
Day 5. By d 5, there was no difference between M and L
or between the M and G time curves. The G and L curves were
not different when the foals were 5 d old (Figures 3.4-3.5).
The S curve also was not different from the other three
curves.

101
Table 3.7. Least squares means for each treatment and each
treatment-day in mmol/liter of plasma glucose.
Treatment
Day 1
Day 3
Day 5
Mean
SE
Glucose
3.98
5.90
5.78
5.22
.25
Lactose
6.44
7.59
7.51
7.18
.25
Maltose
4.57
6.62
6.36
5.85
.23
Sucrose
3.44
5.83
6.29
5.19
.25
Least squares
means were
determined
using a
reduced
model.

Plasma Glucose (mM/L)
Figure 3.1. Changes in plasma glucose over time in 1-day old
foals fed either glucose (SE mean = .15), lactose
(SE mean = .18), maltose (SE mean = .12), or
sucrose (SE mean = .17)
102

Plasma Glucosa (mM/L)
103
Timo aftsr dosing (mln)
Figure 3.2. Changes in plasma glucose (least squares means)
over time in 1-day old foals fed either glucose,
maltose, lactose, or sucrose.

Plasma Glucose (mM/L)
, Changes in plasma glucose over time in 3-day old
foals fed either glucose (SE mean = .26), lactose
(SE mean = .12), maltose (SE mean = .08), or
sucrose (SE mean = .08)
Figure 3.3
104

Plasma Glucose (mM/L)
105
Figure 3.4. Changes in plasma glucose (least squares means)
over time in 3-day old foals fed either glucose,
maltose, lactose, or sucrose.

Plasma Glucose (mM/L)
Figure 3.5. Changes in plasma glucose over time in 5-day old
foals fed either glucose (SE mean = .21), lactose
(SE mean = .13), maltose (SE mean = .13), or
sucrose (SE mean = .11)
106

Plasma Glucose (mM/L)
107
Figure 3.6. Changes in plasma glucose (least squares means)
over time in 5-day old foals fed either glucose,
maltose, lactose, or sucrose.

108
Discussion
Fasting Blood Glucose in Neonatal Foals
The glucose concentration in the blood of very young
foals is higher than in the blood of older foals and adult
horses (Becht and Semrad, 1985). The normal, resting blood
glucose in the adult horse has been reported to be 4.77 ±
.61 mM (Roberts, 1975). Kitchen and Rossdale (1975)
reported glucose concentrations of 5.3 ± .97 mM within the
first 5 min after birth and 4.6 ± .18 mM after 30 min in
seven Thoroughbred foals. All of the foals had nursed by 1
to 2 h postpartum. At 14 h of age, these same foals had
increased their blood glucose concentrations to 7.3 ± .69
mM. Glucose concentrations for foals from birth to 12 h old
range from 6.4 ± .8 to 8.0 ± 1.6 mM (Rose et al., 1979;
Bauer et al., 1984), respectively. Bauer et al. (1984)
determined serum glucose in foals to be 9.2 ± 1.6 at 1 d of
age, 9.3 ± 2.0 at 3 d and 9.0 ± 1.0 mM by 7 d. Rose et al.
(1979) reported somewhat lower values. Foals sampled had
concentrations of 8.0 ± .6 at 12 to 36 h and
8.4 ± .4 mM during wk 1 to 4. Many factors could be
responsible for the difference values reported. One
important factor is time after nursing. None of the workers
noted the elapsed time from the last nursing episode to
sample collection. Even though foals nurse frequently, this
may affect reported concentrations. The foals used in this

109
experiment were fasted for 2 h starting at 6 h postpartum,
before the treatment solution was administered (time = 0) .
On d 1, the 2 h fast caused a considerable decrease in
blood glucose. Even though the mean was 4.14 ± .27 mM, six
of the 13 foals had values of 3 mM or below. By d 3, the
foals were able to maintain higher fasting glucose
concentrations. The foals may have become better at
regulating glucose via insulin/glucagon interactions, more
efficient at gluconeogenesis or better able to spare
glycogen stores with increased utilization of lipid stores.
Perhaps some or all of these systems are not fully
functional at birth. All of the foals were born on or
within a few days of the calculated foaling date, so
prematurity could not be a factor when evaluating the effect
of fasting.
The mean fasting blood glucose concentrations for d 3
(6.08 mM) and d 5 (6.31 mM) were higher than the reported
resting concentrations for adult horses (4.77 ± .61 mM) by
Roberts (1975). This is explained by the major energy
sources utilized in each stage of a horse's life.
Lactose, absorbed as glucose and galactose from the
small intestine, is the major energy source for the very
young foal. As the foal matures, an anaerobic microbial
population becomes established in the cecum and large
intestine. As in the rumen, the fermentation process
produces volatile fatty acids (VFA). These are absorbed and

110
are an important source of energy, along with glucose from
carbohydrate digestion in the small intestine.
Consequently, the adult horse has a higher resting blood
glucose than ruminants which rely heavily on VFA for energy,
and lower than nonruminants which rely mostly on small
intestinal absorption of glucose. For example, normal
resting blood glucose in cattle is approximately 2.78 mM
(Bergman, 1970), while the value for adult swine is reported
to be 5.56 mM (Tumbleson and Schmidt, 1986).
Treatment Curves Pooled Over All Days
The purpose of this trial was to determine the extent
to which very young foals would be able to digest
carbohydrates that may be included in neonatal enteral
formulas. It was presumed that oral glucose would be
absorbed easily and the peaks in blood glucose produced
would provide a reference curve. It also was presumed that
normal, healthy foals would be able to digest lactose. The
resultant curves also would be used for reference. The data
suggested that while oral administration of glucose and
lactose result in increased blood glucose in the young
foals, the pattern of absorption differed. The test for
homogeneity of the two curves shows they are not parallel.
Glucose treatment resulted in short, steep peaks occurring
shortly after time =0. In contrast, oral lactose resulted

Ill
in large peaks occurring later after administration and
remaining high longer than the glucose treatment peaks.
In studies on adult horses, Roberts (1975) found high
peaks in blood glucose associated with galactose
administration. Galactose was only detected in the plasma
with high oral doses. Roberts suggested that in the horse,
galactose is rapidly converted to glucose in the liver, and
then released into the circulation. Very young foals were
not included in Roberts' study. However, data from the
present trial would suggest this also may be the case in
neonates. Future investigation of disaccharide digestion in
the foal might compare the results of lactose, glucose, and
galactose tolerance tests.
It was not known prior to the trial if neonatal foals
had measurable sucrase or maltase activities. The
activities of these enzymes in newborn intestinal mucosa
varies with species. For example, human infants are capable
of digesting both of these disaccharides at birth (Paige and
Bayless, 1987), but calves and piglets are not (Code, 1968;
Manners and Stevens, 1972; Ahrene et al., 1969).
The sucrose tolerance curves remained almost flat
throughout the trial suggesting that sucrase activity was
extremely low. The M curve produced from from data taken
over all 3 d was not parallel to the pooled S curve. The
differences in the treatment curves pooled over all days

112
suggests that there may have been some hydrolysis of maltose
during the trial.
To further investigate the differences in absorption
between the treatments, it is helpful to compare treatment-
day curves.
Day 1 Treatment Curves
On the first day of extra-uterine life, oral glucose
and lactose administration resulted in peaks in plasma
glucose. However, the two curves were not parallel. The
lag time before the peak in the L curves may be due to the
time required for hydrolysis of the disaccharide, or the
hepatic conversion of galactose to glucose, which requires a
series of four reactions.
The S curve decreased slightly after time = 0,
remaining almost flat during the trial period. The M curve,
rising only slightly during the 4 h, was different from the
S curve. It was also different from the glucose and lactose
absorption patterns. The d 1 data suggest that oral
glucose and lactose solutions are absorbed readily, but
sucrose and maltose solutions are not.
Day 3 Treatment Curves
The L and G curves d 3 curves are similar to the d 1
curves, except that the peaks occur earlier in the post¬
administration period. Perhaps time and colostrum ingestion

113
have resulted in increased digestion and absorption
efficiency. In piglets, puppies, and infants, colostrum
intake has a profound effect on the digestive tract,
resulting in increased length, weight, and absorptive
surface area, as well as stimulating the development of
enzymes (Goldstein, 1985; Widdowson, 1985; Widdowson, 1984).
Three-day old foals seem better able to withstand a
short fast, with less decline in blood glucose, than 1-d old
foals. As suggested earlier, perhaps the liver is now more
efficiently using gluconeogenic pathways or mobilizing
lipids to spare glycogen. Insulin and glucagon secretion
may be more finely tuned to maintaining blood glucose
concentrations than they were at birth. The older foals may
have increased glycogen and fat reserves. Further
investigation will be necessary to determine which
mechanisms are involved.
On d 3, the M curve rose slightly during the first
hour. The curve was not demonstrated to be statistically
different from the G curve. This may be due to the
fluctuations seen in the right hand portion of the G curve,
although a characteristic glucose-treatment peak occurred
during the first 30 min. The reason for occurrence of these
fluctuations is not known. Glucose determinations were
repeated on these samples to confirm the results.
On d 3, the M curve was not parallel to the L curve.
The large peak produced from the lactose-treatment data does

114
not occur in the M curve. Therefore, maltase activity in
the small intestine of these foals was extremely low.
Day 5 Treatment Curves
Although the plasma glucose curve after oral lactose
administration contains a large peak during the first 90 min
after treatment and the G curve shows a smaller peak of
short duration, the curves were not statistically different
(P > .05). The M curve, almost intermediate to the G and L
curves especially during the first 90 min, could not be
shown to be statistically different from either of the two
curves. The S curve appears similar to the d 1 and d 3
curves. The sucrose tolerance curve, although almost flat,
was not statistically different from any of the pooled G, L,
or M curves (P > .05).
On d 5, the changes in plasma glucose after oral
glucose and lactose administration are similar to those
observed on d 1. It is not surprising that lactase is very
active in the newborn and that the products of hydrolysis,
glucose and galactose, are readily absorbed. It has been
suggested that in the horse, the rate of galactose
absorption may exceed that of glucose (Roberts, 1975). This
trial was not designed to determine absorption kinetics.
However, on all days the glucose-treatment peak appeared
sooner after dosing than the peak in blood glucose following
lactose treatment. It is not known wether this lag period

115
is due to increased absorption time or to hepatic conversion
of galactose to glucose.
Sucrose does not appear to have been absorbed during
this trial. This agrees with Roberts et al.(1974), who
reported very low sucrase activity in the small intestine of
newborn foals. The activity increased during the first
year, reaching adult concentrations by 7 mo of age. In the
present study, dosing with sucrose on d 1 and 3 did not
result in an increase in sucrase activity on d 5. Dollar
and Porter (1957) could not detect sucrase activity in
neonatal calves, even with sucrose feeding. Sucrase
activity does not appear in calves younger than 44 d old
(Huber et al., 1961). Although piglets utilize glucose and
lactose equally well at birth (Dollar et al., 1957; Ahrene
et al., 1969), sucrase activity does not occur until after
the first week (Huber et al., 1969).
On d 1, maltose administration resulted in a slight
rise in plasma glucose concentration, but no peak was
observed. On d 3 and 5, a slight peak was observed. It is
possible that maltase becomes active near the end of the
first week. Roberts and coworkers (1974) obtained equine
intestinal tracts from slaughterhouses. They detected only
low maltase activity in fetuses close to the end of
gestation. The activity increased during the first month
after birth. No data were given for the very early neonatal
period.

116
Other species also have limited maltase activity
shortly after birth. Piglets are not able to digest maltose
until the end of the first week (Cunningham and Brisson,
1957). Maltase activity could not be stimulated in the
intestine of calves during the first 44 d (Huber et al.,
1961).
Because it was shown in calves (Dollar and Porter,
1957) and foals (Roberts, 1975) that the results of oral
tolerance tests correlate well with tissue analysis for
determining relative enzyme activities, some conclusions can
be drawn concerning maltase, lactase, and sucrase activities
in healthy foals. Lactose, the normal substrate presented
to the intestinal mucosa in newborn foals, was well digested
and absorbed. Sustained peaks in plasma glucose were
produced from the data for each day, shortly after the
treatment was administered. It is likely that other
suitable substrates would have an absorption pattern similar
to that of lactose or glucose.
Low maltose digestion and the lack of sucrose
digestion, even on d 5 of the trial, was evidenced by the
lack of substantial peaks in the plasma glucose curves.
This suggests that maltase and sucrase activities are not
developed sufficiently in foals of this age to support the
use of these sugars in enteral formulas for neonatal foals
less than 1 wk old. Because the inclusion of glucose or
other monosaccharides increase the osmolarity of the diet,

117
they must be used with caution. The choice of a
carbohydrate source is a difficult, but important decision.
Therefore, to discover an ideal energy source for
compromised neonates, other carbohydrates need to be
investigated in future studies.

CHAPTER IV
CONCLUSIONS
The use of animal models for investigation of diseases
in humans has been evaluated and debated extensively.
Realistically, an alternative to performing invasive studies
with humans, especially infants, is needed to provide basic
information. While no model is perfect, it can be argued
that swine are similar to their human counterparts in many
respects. The colostrum-deprived neonatal piglet model
provides many opportunities for basic research in nutrition.
The model may also be used to substitute for foals for many
of the same reasons. Piglets are small, tractable, and
adaptable to laboratory conditions. As a result of this
study, techniques imployed to perform surgery and care for
the piglets were improved and can be used in future trials.
The low survival rate of the Group E piglets fed the
elemental diet did not allow comparison of the two lipid
sources. The piglets apparently lacked sufficient maltase
activity to digest the carbohydrate source of the diet.
Published data, as well as information obtained from this
study, suggest that maltase activity is low compared with
lactase, during the first week of life. Isoflurane
118

119
anesthetia greatly improved the post-surgerical recovery of
the piglets in comparison with ketamine anesthesia, and is
recommended for future studies.
Techniques for mixing formulas, performing surgery, and
raising colostrum-deprived piglets were improved during the
course of this study, and should benefit future trials.
Newborn, as well as 1 wk old sow-fed piglets, had
numerically higher lactase activitiesin the small intestine
compared with sucrase and maltase activities. Sow-fed and
Group C piglets that did not ingest maltose had detectable
maltase activity in the small intestine. Similarly, the
Group E piglets that had not ingested lactose had detectable
lactase activity on d 7.
The piglets kept on trial for 7 d had longer and
proportionally heavier small intestines than did piglets
killed at birth. This is probably due to the effects of age
and feeding.
It was not clear from published reports if very young
foals had adequate maltase activity. Because the
development of disaccharidases before and after birth varies
with species, it was not possible to predict the activities
of these enzymes in foals during the first few days
postpartum. Before engaging in a feeding trial using the
Peptamen formula on foals, it was necessary to determine
their ability to digest this maltose.

120
The results of the foal study suggested that foals 3 d
old and older have an improved ability over newborn foals to
maintain normal blood glucose concentrations after a short
fast.
Foals as young as 1 d postpartum are able to absorb
glucose and digest lactose. Maltose is digested only
slightly by foals on d 3 and 5. The maltase enzyme becomes
active too late in life for a maltose-containing formula to
be fed shortly after birth. Sucrase was not digested to any
appreciable degree even in 5-d old foals. On the basis of
the oral tolerance tests given to foals up to 5 d old,
maltose would not be a suitable carbohydrate source for
inclusion in an enteral formula. Sucrose would not appear
to be a suitable substitute. Glucose was readily absorbed,
but large amounts of this monosaccharide in formulas greatly
increases the osmolarity. This may not be tolerated well by
compromised neonates. The results of this foal trial
indicate that glucose from lactose administration was well
absorbed. However, prematurity or damage to the intestines
may result in reduced lactase activity in neonates.
Therefore, alternative carbohydrate sources appropriate for
compromised neonates need to be investigated in future
trials.

APPENDIX A
PIGLET AND FOAL DATA

Sucrase activity in proximal, middle, and distal sections of
the small intestine of piglets fed an elemental diet for 7 days
Sucrase Activity
/LtMoles glucose produced/hour/gram wet tissue
Piglet
Date of
Birth
TI
MI
BI
P172
Mar 2
0
0
0
P173
Mar 2
0
0
0
P218
Sep 1
3.5
0
0
P222
Nov 3 0
2.2
.8
.4
P226
Nov 3 0
. 6
2.4
1.3
122

Maltase activity in proximal, middle, and distal sections of
the small intestine of piglets fed an elemental diet for 7 days
Maltase Activity
/¿Moles glucose produced/hour/gram wet tissue
Piglet
Date of
Birth
TI
MI
BI
P172
Mar
2
43.0
53.4
156.9
P17 3
Mar
2
107.2
43.6
17.2
P218
Sep
1
25.7
59.6
31.4
P222
Nov
30
35.3
64.9
53.3
P226
Nov
30
21.0
35.2
23.4
123

Lactase activity in proximal, middle, and distal sections of
the small intestine of piglets fed an elemental diet for 7 days
Lactase Activity
/¿Moles glucose produced/hour/gram wet tissue
Piglet Date of TI
Birth
MI
BI
P172
Mar
2
104.0
189.2
555.7
P173
Mar
2
310.3
163.8
133.4
P218
Sep
1
90.7
238.9
90.2
P222
Nov
30
102.7
231.1
187.7
P226
Nov
30
47.5
47.4
37.8
124

125
DATA FROM PRELIMINARY FOAL TRIAL
DAY
1
DAY
5a
TIME
PLASMA GLUCOSE
TIME
PLASMA GLUCOSE
(mins)
(mM/L)
(mins)
(mM/L)
0
7.18
0
8.71
15
8.36
50
8.37
30
4.29
65
7.91
45
3.30
80
7.37
60
6.62
95
7.08
90
2.16
125
6.18
120
4.49
155
6.97
a
On day 5, problems with the catheter prevented the
collection of blood samples at the prescribed time.
Therefore, the times listed are not the same as day 1
collection times.

126
LEAST SQUARES MEANS OF PLASMA GLUCOSE OF FOALS RECEIVING
ORAL GLUCOSE, LACTOSE, MALTOSE, OR SUCROSE ON DAY 1
POSTPARTUM
TIME Lactose Glucose Maltose Sucrose
(mins)
(mM/L)
0
4.77
3.28
3.77
4.81
15
5.34
5.12
4.19
3.38
30
6.34
6.34
4.37
3.43
45
6.99
6.41
5.03
3.40
60
7.78
5.82
5.38
3.36
90
7.99
3.89
5.41
3.32
120
6.92
3.39
5.30
3.35
150
5.70
2.65
5.14
2.80
180
4.97
3.28
5.20
2.99
210
4.68
2.41
5.20
3.23
240
5.11
2.82
5.29
3.12

127
LEAST SQUARES MEANS OF PLASMA GLUCOSE OF FOALS RECEIVING ORAL
GLUCOSE, LACTOSE, MALTOSE, OR SUCROSE ON DAY 3 POSTPARTUM
TIME Lactose Glucose Maltose Sucrose
(min) (mM/L)
0
6.40
5.65
6.24
5.97
15
8.01
7.39
7.16
5.91
30
8.76
6.21
7.20
6.40
45
9.11
5.71
7.11
5.88
60
8.18
5.16
6.79
6.24
90
7.02
5.72
6.71
5.49
120
6.70
6.58
6.34
5.28
150
6.56
5.92
6.49
5.89
180
6.49
5.12
6.88
5.65
210
6.11
5.78
6.50
5.50
240
6.35
5.65
6.42
5.47

128
LEAST SQUARES MEANS OF PLASMA GLUCOSE OF FOALS RECEIVING ORAL
GLUCOSE, LACTOSE, MALTOSE, OR SUCROSE ON DAY 5 POSTPARTUM
TIME Lactose Glucose Maltose Sucrose
(min)
(mM/L)
0
6.63
6.43
5.95
6.44
15
8.58
7.14
7.12
6.77
30
9.39
7.36
7.60
6.83
45
9.10
6.97
7.56
6.59
60
8.49
5.61
7.38
6.21
90
8.07
5.31
6.09
6.34
120
6.21
5.01
6.31
5.97
150
5.81
5.36
5.19
5.70
180
6.56
4.96
5.27
5.69
210
6.25
5.35
5.00
5.46
240
6.23
5.42
5.58
5.87

APPENDIX B
DISACCHARIDASE ASSAY PROCEDURE

130
PROCEDURE FOR DETERMINATION OF LACTASE, SUCRASE, AND MALTASE ACTVITY
IN INTESTINAL TISSUE
This assay is designed to measure disaccharidase
activity in intestinal tissue by measuring the amount of
glucose produced when the tissue is incubated with different
disaccharide substrates. Maltose, sucrose, and lactose are
the substrates used in this trial. When one molecule of
these sugars is cleaved by the appropriate enzyme, one
molecule of glucose is liberated. The exception is maltose,
which releases 2 glucose units. Therefore, the amount of
glucose produced by the maltose incubation must be halved to
calculate the true activity present. After incubation with
the substrate for 30 minutes, the Trinder reagent is added
and incubation continues for an addtional 30 minutes. The
Trinder causes the glucose to be oxidized to gluconic acid
and peroxide. The peroxide then reacts with
4-Aminoantipyrine and p-Hydroxybenzene sulfonate and the
peroxidase enzyme to form a colored compound (Quinoneimine
Dye). The color intensity, which is read at an O.D. of
505nm, is proportional to the amount of glucose produced.
The amount of enzyme present intestinal tissue varies with
species and age. Therefore, the amount of tissue homogenate
to be assayed has to be determined for each type of tissue
analysed. This is done by assaying the tissue using
different concentrations of homogenate in the incubation
tubes. The amount is varied until the amount of glucose

131
produced is within the values of the standard curve and in
detectable amounts.
Preparatory work:
1. Allow spectophotometer lamp to warm up, if necessary.
2. Heat water bath to 37°C.
3. Place Trinder reagent in a beaker and dH20 in a beaker
and place in a covered bucket of ice next to the bath. This
allows time for the Trinder undissolved solids to settle and
for the water to become cold.
Equipment needed:
1. 12x75 mm tubes for incubations.
2. 13x100mm tubes for substrate and sodium phosphate buffer.
3. 9" Pasteur pipettes with rubber bulb for homoginization
procedure.
4. Pipettes.
Preparation of Homogenates from piglet tissue:
Immediately after a piglet was euthanized, the
intestine was removed and carefully dissected from the
mesentary. The intestine was cut into 3 segments of equal
length. The total weight of the empty intestine was
obtained. The proximal, middle, and distal thirds were
frozen and later assayed separately.

132
To reduce sampling errors, the segment of intestine to
be assayed was cut into small pieces, from which random
samples were taken. This was done while the tissue was
partially thawed, but kept cold. A 1 gram sample produces
enough homogenate to perform the disaccharidase, protein,
and DNA assays. A 1:5 w/v homogenate is prepared by adding
5 ml of the homogenizing buffer to a 1 gram tissue sample.
Tissues were homogenized thoroughly in a hand glass on glass
homogenizer, keeping the tissues, buffers, and homogenizers
cold at all times.
The sodium-phosphate buffer and substrates were
pipetted into the test tubes prior to adding the homogenate.
The tubes were incubated immediately. In this trial,
duplicates of each substrate-tissue combination were assayed
with 2 different amounts of homogenate.
Assay procedure:
Several preliminary assays were done to determine the
relative amounts of each of the enzymes present, This was
done to determine the amount of homogenate to be added to
the incubation tubes that would result in detectable O.D.'s
within the standard curve.
Each assay should include reagent blanks, containing
only reagents and no homogenate, and tissue blanks,
containing only buffer and homogenate, but no substrate.
1. Added to each tube:

133
*the appropriate amount of homogenate (according to the
level of enzyme activity present) and .05M sodium phosphate
buffer to total 375 nl.
*300 nl of a substrate (except for "tissue blanks").
Buffer is added to tissue blank tubes in place of substrate
to determine the amount of glucose present in the sample.
2. Vortex tubes. Incubate tubes at 37°C for 30 minutes.
3. Add 375 /¿I cold Trinder reagent. Incubate 30 minutes at
37°C.
4. Add 750 ¿¿1 of cold water to stop the reaction.
5. Read tubes at O.D. 505nm on spectrophotometer. Using a
batch sampler facilitates reading tubes in a short period of
time after the addition of the water. Color is stable for at
least 20 minutes.
Calculations:
Standard Curve;
1. Calculate linear regression line for standard curve, y =
mx + b where y is the absorbance, m is the slope, x is the
glucose concentration, and b is the y-intercept. Use the
linear regression coefficients to calculate the amount of
glucose produced in the sample tubes.
2. Average replicated sample O.D.'s. Subtract the reagent
blank O.D. from these average O.D.'s. The glucose in each
tissue is then calculated using the regression equation
determined by the standard curve.

134
3. The substrates maltose, sucrose, and lactose may contain
free glucose. The tissue samples will also have some free
glucose. The amount of glucose from these sources must be
calculated and subtracted from the amount of glucose
produced by disaccharidase hydrolysis. The tissue blanks
run with the sample (no substrate added) allow for the
determination of glucose in the sample of intestine.
Subtract the calculated glucose in the tissue blanks, the
reagent blanks, and the standard curve blank from the
calculated glucose in the sample. The result is the glucose
produced by enzyme hydrolysis (corrected gluocse). 4.
Divide the corrected glucose (/¿g) by the amount (¿xl) of
homogenate used. If different concentrations of homogenate
were used, average the values obtained from each set.
5. For maltase activity, divide the glucose concentration
(Liq/Hl) by 2 for all tubes containing maltose. The units
are /¿g glucose formed/hr//xl homogenate.
6. /xg glucose//xl homogenate = mg glucose/ml homogenate
Y /xM glucose/mg protein/hr =
X Mg glucose//xl homogenate *
(1-i-. 180) (1/g tissue/g homog) (l/mg_prgtein)
g tissue
(The formula weight of glucose is 180 g/mole.)
The results can be expressed as mM glucose produced/hr/gram
tissue or on the basis of amount of protein or DNA in the
tissue.

135
Prepartion of Buffers
0.5M stock solution of NaH2PC>4 .H^O (monobasic) FW=137.99
Dissolve 68.995g in deionized water (dH20). Bring
volume to 1 liter.
0.5M stock solution of Na2HP04 (dibasic) FW=141.96
Dissolve 70.983g in dH20. Bring volume to 1 liter.
Homogenization buffer (0.01M sodium phosphate buffer, pH
6.0, .002% Triton X-100)
0.01M dibasic solution
Mix 20 ml of the 0.5M dibasic stock solution with
980 ml dH20
0.01M monobasic solution
Mix 20 ml of the 0.5M monobasic stock solution
with 980 ml dH20
Adjust the pH of the monobasic solution to pH 6.0
with the dibasic solution using a pH meter.
0.05M sodium phosphate buffer pH 6.0 (need a minimum of 400
ml to prepare disaccharide solutions)
0.05M dibasic solution
Mix 100 ml of the 0.5M dibasic stock solution with
900 ml dH20
0.05M monobasic solution
Mix 100 ml of the 0.5M monobasic stock solution
with 900 ml dH20
Adjust the pH of the monobasic solution to pH 6.0 with
the dibasic solution using a pH meter.

136
0.5M sodium phosphate buffer pH 7.0
Adjust the pH of 500 ml of the monobasic stock solution
to pH 7.0 with the dibasic stock solution using a pH
meter.
1.0M Trizma base stock solution
Dissolve 60.55g Trizma in dH20. Bring volume to 500
ml.
1.0M Trizma hydrochloride stock solution
Dissolve 78.8g Trizma HCl in dH20. Bring volume to 500
ml.
1.0M Tris buffer pH 7.0
Adjust the pH of 200 ml of the Trizma HCl stock
solution to pH 7.0 with the stock Trizma base
solution.
Preparation of Substrates
0.188M lactose in 0.05M sodium phosphate buffer pH 6.0, a-
lactose monohydrate, crystalline FW=360.32
Dissolve 6.774g in 0.05M sodium phosphate buffer.
Bring final volume to 100ml. Keep on ice or
refrigerated. Do not use after 1 week.
0.0156M maltose in 0.05M sodium phosphate buffer pH 6.0,
maltose, crystalline hydrate FW=360.32
Dissolve 0.562g in 0.5M sodium phosphate buffer. Bring
final volume to 100 ml. Keep on ice or refrigerated.

137
Do not use after 1 week.
0.375M sucrose in 0.05M sodium phosphate buffer pH 6.0,
sucrose, crystalline FW=342.30
Dissolve 1.284g in 0.05M sodium phosphate buffer.
Bring final volume to 100 ml. Keep on ice or
refrigerated. Do not use after 1 week.
Preparation of Trinder Reagent
Combine 50 ml of 1M Tris buffer pH 7.0 and 50 ml of
0.5M sodium phosphate buffer. Add to this the contents of 1
vial of Trinder reagent (Sigma). Stir for 30 minutes.
Allow solution to settle. Use supernatant only. This
reagent is light and temperature sensitive. Store in a dark
container on ice or in cold room.

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BIOGRAPHICAL SKETCH
Lori Pamela Rice was born March 13, 1957, in Cleveland,
Ohio. She lived and attended school in Lyndhurst, Ohio,
graduating from Charles F. Brush High School in 1975. She
then entered the zoology program at The Ohio State
University, as a direct-admission honors student, receiving
a B.S. in 1979 with a combined animal science-zoology major.
During her undergraduate program, she worked at various
veterinary clinics involving large and small animal
practices and became a registered animal health technician.
After graduation, she accepted a position as head laboratory
technician with a large pediatric practice.
In 1981, after having taken several courses locally at
Cleveland State University, she returned to The Ohio State
University. She completed the reguirements for an M.S.
degree in animal nutrition in 1983. Her master's thesis
involved culturing and identifying types of anaerobic
bacteria from the cecum and colon of ponies fed different
diets.
In 1984, she was accepted as a doctoral student in the
University of Florida College of Agriculture. Her
educational program included equine, ruminant, and human
nutrition, reproductive and general physiology, and
biochemistry. The research program emphasized neonatal
146

147
nutrition, equine nutrition, energy metabolism, and
biochemistry.

I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
Edgar A. Ott, Chairman
Professor of Animal Science
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
/
Science
rofessor of Dairy
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
Sandi Lieb
Associate Professor of Animal
Science

I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the deqree of Doctor of Philosophy.
Associate Processor of Animal
Science
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
Peggy L. Borum
Associate Professor of Food Science
and Human Nutrition
This dissertation was submitted to the Graduate Faculty
of the College of Agriculture and to the Graduate School and
was accepted as partial fulfillment of the requirements for
the degree of Doctor of Philosophy.
December 1989
Dean, Graduate School

UNIVERSITY OF FLORIDA
3 1262 08554 2750




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UNIVERSITY OF FLORIDA
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