Blackwell Science, LtdOxford, UKASJAnimal Science Journal1344-39412004 Blackwell Publishing Asia Pty LtdDecember 2004756491498Review ArticleOstrich gastro-intestinal tractR.

G. COOPER and K. M. MAHROZE

REVIEW ARTICLE
Anatomy and physiology of the gastro-intestinal tract
and growth curves of the ostrich (Struthio camelus)
Ross G. COOPER1 and Khalid M. MAHROZE2
1
Department of Physiology, University of Central England, Birmingham, UK and 2Department of Poultry, Faculty of
Agriculture, Zagazig University, Zagazig, Egypt

ABSTRACT
The ostrich is an important animal in many livestock industries and in the developing world. The digestive system of the
ostrich differs considerably from that of non-ruminant animals or poultry. The genetic potential of an ostrich can be
described by its characteristic growth curve. This present paper presents an account of the rearing, anatomy of the gastro-
intestinal tract, and growth curves of this bird.

KEYWORDS: anatomy, gastro-intestinal tract, growth, ostrich, physiology.

INTRODUCTION The ostrich lacks teeth and a crop, and the esophagus
empties directly into the glandular stomach (proven-
The ostrich (Struthio camelus var. domesticus) is the larg- triculus) without any distinct demarcation (Smith &
est of the flightless birds. It is mainly herbivorous and, Sales 1995). Iji et al. (2003) documented an 11-fold
in natural conditions, its diet consists of a variety of increase (P < 0.001) in bodyweight between 3 and
plants and succulents (Jamroz 2000). In free ranging 72 days of age. The relative weight compared with the
conditions, ostriches prefer soft ephemerals to peren- bodyweight of the proventriculus/gizzard, caeca and
nial grasses and shrubs (Williams et al. 1993). High colon increased significantly with age. The relative
mobility and selective foraging are adaptations in arid weight of the small intestine peaked at 41 days old,
environments for optimal use of vegetation (Gatimu from whence it declined (Iji et al. 2003). A recent
1996). The use of drought-resistant fodder crops as an study on ostrich GIT used concentrations of 25–100%
alternative food source has been reported in 8-month- w/vol liquid barium sulfate at 7 and 10 mL/kg, which
old ostriches (Nel & Lambrechts 1998). The common- were administered by stomach tube after withhold-
est drought-resistant fodder crops in South Africa ing food for 16 h (Wagner & Kirberger 2003). The
include the Mexican aloe (Agave americana), thornless Wagner and Kirberger (2003) study follows on from a
prickly pear (Opuntia spp.) and saltbush (Atriplex spp.). previous study in which radiographs were used to
Evolution has selectively resulted in unique adapta- produce schematic illustrations of normal radio-
tions of the gastro-intestinal tract (GIT) of this bird to graphic anatomy (Wagner et al. 2001). Structures
handle such food. The use of the ostrich as a livestock that were consistently identified included the
animal necessitates an understanding of its ability to esophagus, proventriculus, ventriculus, duodenum,
use specific nutrient requirements if efficient growth is jejunum, proximal and distal rectum. Because of the
to be realized.

ANATOMY AND PHYSIOLOGY OF Correspondence: Ross G. Cooper, Department of

Physiology, School of Health and Policy Studies, Baker
OSTRICH DIGESTIVE SYSTEM Building, Room 701, University of Central England, Bir-
mingham B42 2SU, UK. (Email: rgcooperuk@yahoo.com)
The digestive system of the ostrich differs considerably Received 28 May 2004; accepted for publication 29 June
from that of non-ruminant animals or poultry (Fig. 1). 2004.

Animal Science Journal (2004) 75, 491–498 491

R. G. COOPER and K. M. MAHROZE
Fig. 1 The digestive system of an adult ostrich (adapted from
Holtzhausen & Kotzé 1990). (A), proventriculus (glandular
stomach); (B), ventriculus (gizzard) containing approximately
1.5 kg of pebbles; (C), small intestine (approx. 6 m); (D), caeca
or appendices (approx. 0.95–1.25 m); (E), colon or large
intestine (approx. 16 m); (F), hindgut.
superimposition of the remainder of the small intes-
tine, individual components were difficult to differen-
tiate. The caeca were inconsistently highlighted and
only for a short time. The ventral pouch of the copro-
deum never filled with contrast medium (Wagner &
Kirberger 2003). In another study, the coprodeal epi-
thelium was studied by electro physiology in the
Ussing chamber, and the anatomy by light microscopy
and scanning electron microscopy (Skadhauge et al.
2003).
Animal Science Journal (2004) 75, 491–498
Table 1 Comparative measurements (inches) of the gastro-
intestinal tract (GIT) of ratites (Shanawany 1996)
GIT Ostrich Rhea Emu
Proventriculus 14 ¥ 32 4¥4¥8 3¥6¥8
Ventriculus 12 ¥ 16 9 ¥ 9 ¥ 17 4.4 ¥ 6.5 ¥ 10
Small intestine 640 140 287
Large intestine
Caecum 4 ¥ 94 3 ¥ 48 2¥9
Rectum 6 ¥ 800 3 ¥ 40 3 ¥ 25
Liver weight (kg) 2.2 – –
Gallbladder Absent 1.5 2¥8
The ostrich coprodeum is not adapted like a mam-
malian bladder. The abundance of goblet cells results
in a copious secretion of mucus that establishes a thick
unstirred layer giving effective osmotic protection
(Skadhauge et al. 2003). Ostriches like all birds, are
monogastric and their gullet widens to become the
glandular stomach (proventriculus) that joins the
gizzard. The esophagus is markedly wide and when
contracted contains numerous longitudinal rugae
(Shanawany 1996). Comparative measurements (cm)
of the gastro-intestinal tract of some ratites are shown
in Table 1. The esophagus enters the stomach (proven-
triculus). The proventriculus of the ostrich is a large,
dilated thin-walled structure. In ostriches, enzyme and
acid secretion is restricted to an area of only 25% of
the total inner surface area of the proventriculus. This
is in contrast to other birds in which the entire surface
of the proventriculus secretes digestive enzymes. The
distal end of the proventriculus passes to the gizzard
(ventriculus). The ventriculus is a thick-walled struc-
ture. The dark, tough lining (koilin) of the proventric-
ulus and ventriculus are formed by protein secreted
from the glands combined with entrapped sloughed
cells and other cellular debris. The green-brown color
is attributed to the refluxed bile pigments from the
duodenum. The small intestine is relatively short and
straight while the rectum is long and large to enable
the digestion of bulky food and to facilitate fluid
absorption (Shanawany 1996). The round gizzard is
muscular and contains approximately 1.5 kg of rough
pebbles the size of marbles that grind the food as the
muscular walls of the organ contract and relax. The
digestive tract contains two large caecae and a long
colon up to 16-m long (Holtzhausen & Kotzé 1990).
The well-developed hindgut is used as a fermentation
site for plant fiber. The large intestine is three times as
long as the small intestine, although the caeca is of a
similar size in poultry. A spiral fold in the caecum of
the ostrich is described (Bezuidenhout 1993). The fold
492
Ostrich gastro-intestinal tract
spiralled 30 times transforming the lumen into a long,
spiral-shaped cavity. The fold consisted of mucosa,
muscularis mucosa and submucosa. The proximal part
of the fold contained a central core of muscle originat-
ing from the inner circular layer of the caecal wall. The
distal end was expanded and frilled so increasing the
surface area of the fold. The surface area of the spiral
fold averaged 955.75 cm2 constituting 54% of the total
caecal mucosal surface.
Bezuidenhout (1993) suggested that the fold plays
an important role in the absorption of volatile fatty
acids and other metabolites produced by microbial fer-
mentation of cellulose and hemicellulose. Although
there are many similarities between ostriches and
chickens, differences include simple superficial
proventricular glands, branched tubular glands, and
deep proventricular glands restricted to a slipper-
shaped arc and extending into the muscularis mucosa
(Bezuidenhout & van Aswegen 1990). A unique fea-
ture in the ostrich is the variably developed muscularis
mucosae in the gizzard. The villi of the small intestine
are long and branched profusely forming a labyrin-
thine surface. No Paneth cells were observed. The
presence of numerous melanocytes in the submucosa
and connective tissue around the blood vessels of the
muscle layers at the tips of the caeca is surprising, as
these cells are normally associated with pigmentation.
A well-developed subserosa was observed throughout
the gut. The topographical distribution of endocrine
cells immunoreactive to glucagon, bombesin, neuro-
tensin and substance P differed from that of the
chicken. Secretin was structurally different from its
counterparts in the mammal and chicken (Bezuiden-
hout & van Aswegen 1990).
The topography of the thoraco-abdominal viscera
in 20 ostriches aged 2 weeks to 12 months has been
described (Bezuidenhout 1986). Important anatomi-
cal positions of the gut include the proventriculus
that occupies the left cranial part of the abdomen,
resting on the sternum and abdominal floor. The
duodenum formed a loop from right to left with the
pancreas lying between the two limbs. The coiled
jejunum and ileum were located in the ventral part
of the abdomen between the gizzard and pelvis. The
two caecae were situated on either side of the termi-
nal ileum with their apices in the pelvis. The rec-
tum was the longest part of the intestine and was
divided into a thick proximal segment situated in the
right dorsal part of the abdomen and a thin distal
part that occupied the left caudodorsal part of the
abdomen.
Animal Science Journal (2004) 75, 491–498
EXCRETA FORMATION AND
MOVEMENT IN THE GASTRO-
INTESTINAL TRACT
In one review, Duke (1999) detailed excreta formation
in ostriches as liquid urine followed by discrete faecal
boli excreted in rapid succession. The cloacal structure
of the ostrich possibly explains the type and mecha-
nism of excreta formation. Rectal stricture greatly
impedes excreta formation and elimination. Indeed,
Berridge et al. (1998), in a exploratory celiotomy of an
18-month-old female ostrich with anorexia, lethargy
and constipation of 5 days duration, showed mesen-
teric volvulus and a focal narrowing of the rectum
~28 cm cranial to the cloaca. The animal was eutha-
nized and necropsy revealed a fibronecrotic proctitis
and a 3 cm long circumferential stricture of the rec-
tum. Lower intestinal function was compromised by
the rectal wall at the stricture that was deeply effaced
by fibrovascular connective tissue with vascular
thrombosis and necrotizing vasculitis. The emu, how-
ever, has a limited ability to produce concentrated
urine (Dawson et al. 1985). Eight emus were water
restricted and the osmolality of the excreta fluid was
shown to increase to a maximum after 6 days to
428 ± 39 mmol, the urine:plasma being 1.35 ± 0.12. In
normally hydrated birds most of the Na+ and Cl– ions
were excreted in the fluid fraction although one-third
of the K+ was associated with the solid phase. After
water restriction for 9 days the pattern of K+ excretion
did not change, although the majority of Na+ (79%)
and Cl– (73%) were excreted in the solid phase possi-
bly associated with urates. The cloaca-rectum appears
to play a major role in the modification of excreta. It is
conceivable that a similar physiological function
occurs in the ostrich, although this would require fur-
ther investigation.
Retrograde movement of urine from the coprodeum
to the colon and caeca in birds is important in the
recovery of water and sodium chloride (Duke et al.
1994). Water is derived from both hypotonic luminal
fluid and hypotonic fluid if the rate of Na+-linked
water absorption exceeds the rate of osmotic water
loss. Ostriches (aged 10 months; 90–10 kg bwt) were
slaughtered and used for anatomical examination.
They were raised on desert range (temperature of 32–
34∞C in August; annual rainfall of 200 mm). The birds
were allowed to feed on natural vegetation and were
given commercial feed (2 kg/bird/day) and fresh Luc-
erne. The lower colon and cloaca were collected from
16 ostriches at slaughter. Four initial samples used to
493
R. G. COOPER and K. M. MAHROZE
visually differentiate and determine the coprodeum
and proctodeum, were combined with 12 subsequent
samples to determine if urine passed into the terminal
colon (presence of uric acid) and the distribution of
urine between the coprodeum and proctodeum. Lon-
gitudinal opening of chambers and the presence of
uric acid and urine, and the collection of urine samples
from 10 cloacae were analyzed. Serial radiographic
observations using a suspension of 25% (w/v) barium
sulfate (BaSO4) was administered into the coprodeum
of the adult birds. Urine was stored principally in the
proctodeum that changed in size with urine volume.
In 8-week-old chicks, it was 6–8 cm in diameter prior
to urination when normally hydrated and 1–2 cm fol-
lowing urination. Mean osmolality of the cloacal fluid
was 395.4 mOsm/L (SD = 167.5; range = 185–715;
n = 10), which was ~1.2–1.3 times the plasma osmo-
lality (300–330 mOsm/L) (Withers 1983). The major-
ity of BaSO4 suspension administered into the
coprodeum moved into the proctodeum by 20 min.
Similarly the contrast medium administered to chicks
was cleared from the blood by the kidneys, passed
through the ureters and quickly through the copro-
deum into the proctodeum within 25–40 min. Urine
and feces were excreted separately (urine first) and
the birds were coprophagic.
Degen et al. (1994) investigated the sequence of gas-
troduodenal contractions and the functions in the
stomach in two male ostriches aged seven and eight
weeks. Birds housed in an animal-holding room were
subjected to a relative humidity (RH) of 45–50%, tem-
perature of 25–27∞C and lighted conditions from 06.00
to 20.00 hours daily. BaSO4 was mixed with mash diet
so that movements of food within the gut could be
observed using radiographic imaging. Gastric contrac-
tion frequency increased in one bird from 2.08 ± 0.32
(hydrated) to 2.68 ± 0.32 contractions/min (dehy-
drated), and decreased in the other from 2.08 ± 0.40 to
1.56 ± 0.23 contractions/min. However, no cause for
the different responses between the two birds was
apparent. The duodenal reflux might account for the
oral flow of ingesta from the upper half of the small
intestine into the stomach. This presumably improves
the overall digestibility of the diet by exposing it 2–3
times to duodenal and pancreatic digestive enzymes.
The hydrated ostriches always had a large volume of
water in their glandular stomachs. Water deprivation
for 20 h, however, resulted in stomachs devoid of
water. The authors described this as an indication of
the duration of water storage in the reservoir. In adults
or larger juveniles with a lower metabolic rate and rel-
Animal Science Journal (2004) 75, 491–498
atively less body surface area, the reservoir might last
longer (Degen et al. 1994).
RESISTANCE TO DEHYDRATION
Living in semiarid or arid conditions has favored the
ability of ostriches to withstand substantial dehydra-
tion (up to 30% body mass loss) and thereafter drink
water approximately equating the body mass lost dur-
ing dehydration (Cloudsley-Thompson & Muhamed
1967). The rapid ingestion of a large volume of water
(17% body mass) following dehydration only caused a
slight decline in plasma osmolality (Withers 1983)
suggesting that the water entered the plasma slowly.
Degen et al. (1994) described that ingested water is
stored in the glandular stomach and then released
slowly until the next water ingestion. This allows
ostriches to graze over a large radius from a water
source because their great mobility and they can
replenish the stomach in one ingestion. There has
been no mention of the glandular stomach in other
avian species although it might act as a food store in
emus and the lower small intestine might act as a
region for water storage (Herd & Dawson 1984). The
glandular stomach in ratites achieves the storage func-
tion of the crop in other birds.
GROWTH CURVES FOR OSTRICHES
The growth dynamics of the primary cortical bone of
young ratites shows that from hatching to 2 months of
age, cortical thickness remains constant thus express-
ing equilibrium between periosteal bone deposition
and an endosteal bone reabsorption. Radial growth
rates of the diaphyseal bone cortex are high (10–
40 mm/day on average; maximum of 80 mm/day) in
the hind limb (femur, tibiotarsus and tarsometatar-
sus). Wing bones are smaller and develop later (Cas-
tanet et al. 2000). The genetic potential of an ostrich
can be described by its characteristic growth curve. A
Gompertz model on growth performance of ostriches
from different localities in southern Africa has been
described (Du Preez et al. 1992). The results of Du
Preez and coworkers were verified using 43 mixed sex
birds solely at Oudtshoorn (Cilliers et al. 1995). Body-
weights at hatching were 0.81–0.85 kg for males and
0.78–0.85 for females. Mature weights estimated from
the Gompertz curve, were 94.2–102.1 kg for males
and 92.6–104.6 kg for females. Ages of maximum
daily gain were 92–163 and 114–175 days for the two
sexes, respectively. The estimated mean mature
494
Ostrich gastro-intestinal tract
(14 months) bodyweights were 119.2 kg for males and
122.3 kg for females, parameters that did not differ sig-
nificantly between sexes. There were significant dif-
ferences for most growth parameters among birds
differing in origin from Oudtshoorn, South Africa,
Namibia or Zimbabwe.
The Gompertz growth curves have multiple uses in
production and research and are used as a tool to mea-
sure the standard of management and feeding versus
growth potential of the birds (Smith & Sales 1995). A
method of imitating the feeding pattern of ostriches in
the wild would be useful as it would give a good indi-
cation of nutrient requirements in captive birds. Nutri-
tionists could use different approaches to study the
effects of nutrients on growth patterns (Du Preez
1991). Male ostriches might have to be raised sepa-
rately as they grow faster, require higher protein diets
and are more efficient feed converters for longer
periods than females (Shanawany 1996). Significant
(P < 0.01) differences in the rate of growth between
birds from different regions and sex differences were
demonstrated (Du Preez et al. 1992). Growth curves in
ostriches showed a faster age-gain in weight for birds
in Zimbabwe (Fig. 2). Carcass analysis from birth to
maturity to determine the development of chemical
components of the body during growth has been
described (Du Preez et al. 1992). The sacrificed birds
were defeathered and the empty body (minus gastro-
intestinal tract) was analyzed for moisture, ash, pro-
tein and lipid. Dry, lipid-free samples were also ana-
lyzed for amino acids. Protein accretion in the body
was used as the system to calculate amino-acid
requirements of the birds. Because of the lack of data
on ostriches, poultry coefficients (0.73 and 0.63) were
used, respectively, to convert the dietary amino acids,
lysine and sulfur-containing amino acids into protein
100
80
Weight (kg)
60
40
20
0 (6 ¥ 200 m) and at 4 months (50 ¥ 250 m). At
0 100 200 300 400 500 600
Age (days)
Fig. 2 Fitted growth curves for ostriches in three localities
(source: Du Preez et al. 1992).
Animal Science Journal (2004) 75, 491–498
accretion. The estimated protein requirement for
maintenance was calculated using the following for-
mula (Emmans 1989):
MP = 8Pm0.73u
where MP = protein requirement for maintenance;
Pm = mature body protein mass (kg).
u = P/Pm
where P = body protein mass (kg).
In a study of protein requirements of ostriches (60–
110 kg bwt), in which lysine was 5% of the dietary
protein, the highest gain in weight (235 g/day)
occurred when the birds were consuming ~2.2 kg/day
of a diet containing 14% protein. This equated to 15.2-
g/day lysine. Estimation of daily feed requirements to
meet the energy needs of birds can be determined
using the formula (Emmans 1989):
MH = 1.63Pm0.73u
where MH = maintenance heat (MJ/day); Pm = mature
protein mass (kg),
u = P/Pm
where P = body protein mass (kg).
Estimates of daily gain in protein and lipid were
converted into daily energy requirement. Adding this
to the maintenance requirement allows determination
of the total energy requirement to which energy
expenditure during activity and locomotion are added.
Growth results of 43 ostrich males and females were
used to compile growth parameters of a flock that is
representative of Oudtshoorn birds using the Gomp-
ertz model. In the paper, there was verb confusion
with the use of ‘chickens’ to represent ‘chicks’.
Growth data were derived from 19 recordings of body-
weight for each individual bird from 1-day-old to day
520. Chicks were reared in floor pens in a semi-
Oudtshoorn
environmentally controlled chicken house from 1-
Namibia day-old to 2 months of age. From one week old, chicks
Zimbabwe
were allotted to small outdoor camps (6 ¥ 10 m) on
Lucerne pastures during the day. Ambient tempera-
tures were maintained between 23 and 25∞C with the
use of overhead radiant heaters. At 2 months old, the
birds were moved to larger Lucerne camps (6 ¥ 50 m).
The size of these camps was increased at 3 months
700
5 months, birds were removed from Lucerne pastures
and assigned to two identical feedlots (25 ¥ 25 m).
Male and female chicks were not separated. Six diets
were offered to the birds during the experimental
495
R. G. COOPER and K. M. MAHROZE
period: pre-starter (0–2 months); starter (2–
4 months); grower (4–6 months); finisher (6–
10 months); post-finisher (10–14 months); and
maintenance (14–18 months).
Competent use of statistical analysis was presented
in the implementation of Statistical Analysis Systems
to fit Gompertz curves to the collected data (sex, age
and weight). Each individual data was used to fit a
curve to obtain estimates of maximum weight at
maturity (A) and the rate of growth (B), also referred
to as rate of maturing. The curves were forced to pass
through the mean hatching weights of all individuals,
and A and B were estimated thus applying the same
form of the Gompertz equation used previously (Du
Preez et al. 1992). The time of maximum daily weight
gain was computed using the estimated parameters for
A & B. The various estimates for these parameters
were compared between birds by t-test. Males exhib-
ited slightly faster maturation rates (¥10-3) than
females (mean (SE): 9.10 (0.278) and 8.55 (0.412),
respectively), although this did not differ significantly
from females. The corresponding age at which maxi-
mum growth was attained was 181 days for males and
199 days for females. Generally the age of maximum
growth in animals usually occurs at one-third of the
age before maturity. The inflexion point of curves for
male and female birds showed that the expected ages
of maturity were 602 and 664 days, respectively. As
the birds were only weighed until 520 days, they were
not fully mature at the end of the study. Comparison
of mature bodyweights of cocks and hens showed sig-
nificantly (P < 0.05) lower estimates of 102.1 and
98.4 kg, respectively. No significant differences in mat-
uration rate were, however, observed between the
findings of the present and former studies (Du Preez
et al. 1992). The former demonstrated estimates of
mean (SE) 9.7 (0.43) and 9.0 (0.44) for cocks and
hens, respectively, versus estimates of 9.1 (0.28) and
8.5 (0.41) in the latter. Calculated ages at which max-
imum weight gain occurred were substantially earlier
at 163 and 175 days for cocks and hens as opposed to
181 and 199 days in the present study.
The importance and usefulness of the Gompertz
curve to describe animal growth is evidenced in the Du
Preez et al. (1992) study as a tool to measure the stan-
dard of management and feeding against the potential
growth of the animal. Additionally, it can be used for
statistical comparisons of measured growth character-
istics between birds to help in the selection of progeny
for future breeding. The ability to quantify the effect of
intensity of feeding or nutritional standards on the rate
Animal Science Journal (2004) 75, 491–498
of growth should be considered. The authors suggest
further studies to determine to what extent the restric-
tion of a limiting amino acid influences growth rate.
Such a study should be supported by carcass analysis.
Improvements in grading of ostrich parts using
the Gompertz (1825) function have been described
(Mellett & Randall 1994). The authors noted that in
nutritional and other growth studies that repeated
measurements are made on the same animals as time
progresses, whereas in meat studies the animals are
slaughtered and only one measurement is possible. In
the study, 10 ostriches were slaughtered aged 3, 6, 8,
10, 12, 14, 16 and 18 months, respectively. Following
the determination of the mass of body parts, a Gomp-
ertz function (Rogers et al. 1987) was fitted to these
measurements and the point at which growth
occurred at its maximum rate was computed by equat-
ing the second derivative to zero. Only the growth of
the head (y in g) versus time (x in months) could be
described with the Gompertz function:
613
y = 613 expÈÍ -ln exp( -0.26x )˘˙.
Î 133 ˚
Growth rate, total body water volume (TBW), dry
matter intake (DMI) and water consumption were
determined in ostriches from hatching to 350 days
(100 kg bwt) (Degen et al. 1991). The Gompertz equa-
tion was used to describe the sigmoidal growth curve;
mature body mass (Mmb) was calculated as 104.1 kg.
The highest average daily gain (ADG) was 455 g/day
that occurred between 70 and 98 days. The growth
curve of ostriches was sigmoidal and asymptotic mb
was calculated as 104 kg reached in 1 year. The met-
abolic age of ostriches at hatching was 13.7 days,
which was similar to the 14 days reported for broilers
(Wilson 1977) and lower than the mean of 23.6 days
for six precocial birds (Kirkwood & Webster 1984).
The metabolic age of the ostriches was lower than the
mean of the six birds to reach 0.5 Mmb (48.6 vs mean
of 67.8) but the time required to grow from 0.25 to
0.75 Mmb was longer (39.4 vs mean of 34.8). Total
body water volume (TBW) as a fraction of mb
decreased from 0.84 (35 day olds) to 0.57 (322 day
olds), attributed to the accumulation of lipid. DMI also
followed a sigmoidal curve and plateaued at 2.05 kg/
day. The ratio of DMI to ADG increased from 1.07 (35
to 42 days) to 17.1 (322–350 days). This explanation
forwarded was during growth of the ostrich, its abso-
lute maintenance requirements increased while its
metabolisable energy intake above maintenance
decreased following an initial increase. As ostriches
496
Ostrich gastro-intestinal tract
matured, the fraction of total body solids increased
including an increase in the fraction of body lipid
content. More energy was thus required per unit of
growth (Degen et al. 1991).
CONCLUSION
Accurate evaluation and knowledge of the nutrient
contents of raw materials for formulating diets and
knowledge of nutrient needs during various stages
of growth and development, and production of the
ostrich are vital. More research could be done to
investigate different food sources by country/region
and how this affects growth, development and per-
formance of the bird. Collaborative research will be
of benefit to mankind because of the healthy red
meat and luxurious leather the ostrich provides, in
addition to the many other potential health-benefits
of its body parts, such as corneas for eye
transplants.
ACKNOWLEDGMENTS
The library assistance of Mr D. Swanepoel and Mrs F.
Jones in South Africa is appreciated.
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