RESEARCH PAPERS
Foamy Bloat of Cattle. A Review
INTRODUCTION
Bloat is a disorder of ruminants caused by
retention of gas in the stomach. Gas is pro-
duced continuously and in large amounts in
the ruminant stomach and is eliminated nor-
maUy by the animal without difficulty. Some-
times, however, the rate of elimination falls be-
hind the rate of production so that gas starts
to accumulate in the rumino-reticulum (here-
inafter called the rumen). If the disparity con-
tinues, it can lead to gross distention of the
rumen -- the intraruminal pressure may rise
to as much as 70 mm Hg (41, 89) -- and the
animal becomes bloated. The inflated stomach
mechanically interferes with respiration, which
probably is further embarrassed by the absorp-
tion of CO2 from the rumen, and can cause
death within an hour of commencing to feed
(81, 105, 112, 315, 388).
The imbalance of gas production and re-
moval that leads to bloat results most common-
ly from excessive foaming of the contents of
the rumen; this is the primary cause of bloat
in animals feeding on legumes or lush young
grass, and of feedlot bloat. Non-foamy or free-
gas bloat is less common than foamy bloat and
results from a variety of causes.
This review is concerned with foamy bloat,
and especially legume (pasture) bloat. Feedlot
bloat, which characteristically oeettrs in beef
cattle fed on rations high in concentrates and
low in roughages, and which may be chronic,
will be discussed separately and in less detail.
Extensive reviews of bloat appeared regularly
between 1945 and 1960 (81, 82, 83, 190, 191)
the last being that of Cole and Boda (81) in
1960. Since then the subject has been consid-
ered in varying degrees of detail by a number
of authors (66, 78, 175, 183, 196, 322, 327,
379). The present review, which mainly covers
the literature since 1959, should be read in as-
sociation with that of Cole and Boda (81).
INCIDENCE AND ECONOMICS
Bloat, whether occurring on pasture or in
the feedlot, is a widespread complaint as is
shown by the published literature arising from
Received December 21, 1972.
753
R. T. J. CLARKE and C. S. W. REID
Applied Biochemistry Division
Department of Scientific and Industrial Research
Palmerston North, New Zealand
at least 23 countries. I t is predominantly a dis-
order of cattle but may occur .also in sheep and
other domestic ruminants (57, 399) and has
been observed in wild antelopes (154). There
is evidence for differences between cattle
breeds in their susceptibility to bloat (205,
270, 295, 328). Within breeds there is a wide
range of individual susceptibility from animals
of high susceptibility (HS animals) to animals
of low susceptibility (LS animals) (78, 328).
Young animals have been reported more sus-
ceptible than old (80, 295).
Bloat occurs mainly on legume forages, but
there are also reports of its occurrence on suc-
culent grasses, various crops such as cereals
and brassicas (190), and hay (3). Not all
legumes are bloat-inducing: for example, there
are few reports of bloat when tropical legumes
are fed (164, 178, 209). The species most
commonly causing bloat are alfalfa (Medicago
sativa) and various clovers (Tri[olium spp.).
Unless otherwise stated, the word legumes will
refer to bloat-inducing species.
Bloat is associated with lush growth, and its
incidence generally reflects the seasonal
growth patterns of these legumes: factors fa-
voring legume growth typically increase the
incidence of bloat and vice versa. The use of
mixed pastures tends to reduce the incidence
of the disorder (345, 404, 405) as does the use
of mature rather than immature legumes (165,
295) or the feeding of legumes as hay rather
than green feed; but bloat can occur in all of
these circumstances (295, 318).
The full economic cost of bloat is not easy
to deduce. Losses or costs of several kinds oc-
cur: losses by death and by culling of bloat-
prone animals, losses of production from ani-
mals which bloat and survive, losses due to the
disruption of normal farm work and manage-
ment programs, losses due to the use of less
productive but safer pastures, and the cost of
preventive measures and treatment. Not meas-
urable in monetary terms is the mental and
physical strain on the farmer and his family
when an outbreak occurs. To these costs must
be added those of research and extension serv-
ices.
Death of animals is the most obvious loss
eansed by bloat, and the cost of their replace-
754 CLARKE A N D R E I D
ment is usually the only basis used for estimat-
ing the economic cost of the disorder.
Reductions in milk yield and in growth rate
which typically accompany bloating in cattle
are due basically to reduced food intake. In
stall-fed animals even mild bloat will depress
intake (330) while cows at pasture usually
stop grazing when they become moderately
bloated (165). Hancock (165) found grazing
and (especially) rumination times reduced
more or less in proportion to the extent of
bloating. A fall in milk production is charac-
teristic of a severe outbreak in a dairy herd,
but in milder outbreaks, milk yield may be
little affected (344). In beef cattle reduced
weight gains have been associated with the
grazing of pas~tres with a high content of
clovers because of greater incidence of bloat
on the clover-dominant pastures (6, 405). Cat-
tle, however, will produce more on legume
pastures than on grass ff an effective bloat pre-
ventive is used (1, 2, 366, 368, 369). Bloat al-
so reduces the weight gain of cattle in the
feedlot (147, 270).
The greatest economic loss from bloat prob-
ably occurs in the U.S.A. The U.S. Department
of Agriculture (380) estimated the average an-
nual loss, 1951 to 1960, as $US 105,000,000
for cattle alone. This was about equally divid-
ed between beef and dairy industries. In a sur-
vey of feedlots in Kansas (E. E. Bartley, per-
sonal communication) involving some 720,000
cattle, in 1969 about .1% died of bloat while
a further .2% had bloated severely and .6% to
a lesser degree. Dougherty (78) has referred
to a feedlot raising 120,000 cattle a year in
which about 1% of the animals were seriously
bloated, nearly half of these dying from the
disorder.
In Australia, pasture bloat is of considerable
importance to the dairy and beef industries of
Victoria and New South Wales, but it is not
of significance in the north, nor is it of conse-
quence to the Australian sheep industry (10).
Published figures for mortality from bloat
range up to 10% (286). In one survey, deaths
ranged from .27% in a drought year to 2.5 to
3.0% in a year of average or better rainfall
(404).
In New Zealand bloat is a serious disease of
dairy cattle, but it is not yet a serious problem
in beef cattle (327) or in sheep. The national
loss of dairy cattle by death from bloat for the
seasons 1964 to 71 was .2 to 1.2% (295) with
the mortality in individual herds sometimes ex-
ceeding 10%. The total annual losses from all
aspects of bloat could be $NZ 8,000,000.
In Canada the losses from sickness and
]OURNAL OF DAIRY SCIENCE VOL. 57. N o . 7
death have been estimated by Miltimore
(272) to be SCan. 11,000,000 annually. No
official statistics on the incidence of bloat are
kept in Great Britain, Argentina, South Africa,
or Eire.
Although sheep have often been used as ex-
perimental animals in bloat research, the disor-
der seems of less importance in sheep than in
cattle. We could find no data on the economie
importance of bloat in animals other than cat-
tle.
THE RUMEN GAS POOL AND ERUCTATION
Gas is in the rumen in two interchanging
states, dissolved and free, which together make
up the rumen gas pool. Bloat may be regarded
as a major increase in the pool size due primar-
ily to interference with gas outflow.
Most of the gas in the rumen is formed
there. The main sources are microbial fermen-
tation and acidit~cation of bicarbonate; the ma-
jor components are CO2 (45 to 70%) and CH4
(20 to 30%), with N2, 02, H2, and H2S as mi-
nor components. The daily total gas production
is, theoretically at least, large, but in bloat the
important factors are (1) the rate of produc-
tion during feeding, especially the peak rate
and (2) the degree of gas accumulation. Esti-
mates of the rate of gas production in cattle
range from less than .2 liter/min in the fasted
animal (173, 393) to 2.0 liter/min following
feeding (93, 173, 176, 222, 393). If gas is al-
ready accumulating in the stornaeh, small
amounts of additional gas may increase mark-
edly the severity of bloating (190).
There are three major pathways out of the
gas pool: absorption through the rumen wall,
passage out to the omasum, and passage out
to the esophagus. The last pathway, eructation,
is the most important for free gas, particularly
for gases of low solubility such as CH4 and N2.
The other pathways cannot cope with normal
gas production ff the esophagus is obstructed.
Eructation is a complex event, requiring co-
ordination of the activities of several organs
and structures. It is trader control of the cen-
tral nervous system, and the necessary coordi-
nation is achieved by operation of a series of
nervous reflexes. Discussions of the physiology
of eructation in ruminants are given by Dough-
erty (109, 110, 117), who with his co-workers
has been responsible for elucidating many of
its facets, and by others (81, 179, 323, 334,
359, 376, 398).
The movement of gas during eructation
takes place in several stages: (1) separation
stage in which gas forms as bubbles and rises
through the digesta to coalesce with the free
 REVIEW: F O A M Y
cap in the dorsal rumen; (2) displacement
stage during which the gas cap is moved for-
ward and downward to the cardia; (3) trans-
fer stage during which the cardia opens and
the gas passes out of the rumen into the
esophagus, accompanied hy a sharp contrac-
tion of abdominal wall muscle; (4) esophageal
stage in which, associated with a fast-traveling
contraction of the esophagus, the gas passes
rapidly up the esophagus and out through the
superior cervical esophageal spincter to the
pharynx (114); (5) pharyngeo-pulmonary
stage during which the gas, restrained by clo-
sure of the nasopharyngeal sphincter, passes
through the open epiglottis and is forced into
the lungs where a large portion of the gas is
absorbed, the remainder being exhaled in the
next expiration.
Eructation is usua/ly associated with rumen
contractions, probabl); because the cardia is
normally submerged by digesta which must be
redistributed to allow access by the gas cap.
The most effective for this purpose ,and those
with which eructation is most eommouly asso-
ciated, are the B sequences (the rumen con-
tractions that occur independently of reticulum
contractions) (323, 324, 329, 333, 376), also
called the secondary or forward-moving con-
tractions. In addition, eructation may also oc-
cur with A sequences (those rumen contrac-
tions associated with reticulum contractions)
and, on occasions, without any obvious rumi-
nal movements. An important factor determin-
ing the relationship between eructation and
rumen movements seems to be the degree of
fill of the rumen. This contention is given sup-
port by the behavior of the system when gas
is insufflated into the rumen. Insufflation is ac-
compauied by a fall in the liquid in the lumen
as gas accumulates (115, 323, 360), a simple
physical result of distention. Radi01ogical
examination of chaff-fed sheep during insuffla-
tion after an overnight fast (323) revealed
that the liquid, already low, fell quickly, to ex-
pose the dorsal edge of the anterior pillar and
the eardia, hy which time eructation was oc-
curring freely, associated with B sequences, A
sequences, or without rumen movements. After
the sheep had been fed, however, liquid was
substantially higher and did not fall so readily
during insufflation. Eructation occurred prin-
cipally with B sequences: eructations with A
sequences were less frequent and apparently
smaller while few eructations occurred without
rumen activity. A relationship between fill and
eructation efficiency has been demonstrated
by several workers (107, 233, 287, 398).
Akester and Titchen (4) have described dis-
BLOAT 755
placement of the gas cap during B sequences:
their observations, made radiologically, would
also suggest A sequences to be important in as-
sisting the separation stage.
Opening of the cardia usually occurs in rela-
tion to rumen contractions (359, 398) al-
though it also must open for eructations with-
out rumen movements. Little is known about
the manner of control of the cardia. Stevens
and Sellers (359), in experiments with cattle,
found that eructation occurred only in associa-
tion with rumen movements, and then more
frequently with B than with A sequences, even
though the cardia was kept completely clear
by emptying the rumen or by confining the
contents to a large plastic bag.
During insufflation experiments at this lab-
oratory, cattle have eructated both with and
without ruminal movements. Eructation usual-
ly occurred with the B sequences, but some in-
dividuals were more prone than others to eruc-
rate with A sequences as well (329). As in
sheep, eructation without rumen movements
was uncommon in fed cattle.
Eructation and B sequences have been ob-
tained as separate reflex responses to insuffla-
tion of the rumen in decerebrate sheep (334).
The site of receptors involved remains uncer-
tain. Weiss (398) believes distention of the
caudal dorsal rumen to be the stimulus in-
volved in B sequences while Leek (233) con-
siders the cranial rumen to be the reflexogenic
site. Reid (323), using conscious, chronic
spinal sheep (spinal cord cut between C7 and
T1), has elicited B sequences as a reflex re-
sponse to localized stretch of the rumen at sev-
eral locations, including the posterior pillar. B
sequences were also elicited by gentle rubbing
of the anterior pillar or ventral rumen, an ob-
servation that may be significant in relation to
the improved efficiency of eructation observed
by Colvin et al. (89) when hay was added to
the diet of cattle grazing alfalfa pasture. Reflex
inhibitory responses also have been obtained
(113, 233, 323, 334, 398). Dougherty et al.
(113) have described inhibitory effects of
flooding the eardia with fluid. Inhibition of B
sequences as a response to distention of the
reticulum has been observed by Reid (323),
Reid and Titchen (334), and Leek (233), and
ascribed hy Leek to high threshold receptors.
Manipulations of wounds, or other maneuvers,
which in conscious animals would be expected
to stimulate pain receptors, have caused inhibi-
tion of gastric motility in anaesthetized (233)
and in decerebrate sheep (374, 334). These
several observations must be viewed in the
light that reflex activity will be moderated by
JOURNAL OF DAIRY SCIENCE VOL. 57. NO. 7
756 CLARKE AND R E I D
circulating hormones, metabolites, and electro-
lytes, and by the acid-base status of the animal
(62, 94, 375, 376).
The functioning of the cardia, esophagus,
and pharyngeal structures during eructation,
aspects of the anatomy and innervation of
these structures, and the deposition of gas in
the lungs, have been studied by Dougherty
and co-workers (110, 117). They have also
demonstrated, using :4CO2 and :4CH4, an ex-
tremely rapid absorption of gas, especially
COz, and its incorporation in milk and saliva.
This has led to the investigation of eructation
as a likely means of absorption of taints se-
creted in milk (118, 350).
That eructed gas enters the trachea (88,
119) has been the basis of a method of collect-
ing eructed gas for measurement and analysis.
Gas entering the upper trachea is led away
through a tracheal fistula while respiration con-
tinues through a fistula in the lower trachea,
or a form of cannulation of a single fistula al-
lows a similar separation (88, 119, 387). Hoer-
nicke et al. (173) used this method with x4COz
and :4CH~ to investigate the proportions of
each gas absorbed and eructed. The success
of the method depends on the proportion of
eructed gas that is lost out of the mouth in-
stead of entering the trachea. Dougherty and
Cook (111) found losses were greater in one
cow than another and higher during rumina-
tion; Hoernicke et al. (174) reported losses of
.5 to 33%. The method has not received the
attention it deserves; apart from the initial
work of Colvin et al. (88) it has not been used
since in bloat studies.
FOAMY BLOAT
Introductory
Acute pasture bloat is caused by the devel-
opment of a stable foam in the contents of the
rumen (41, 67, 176, 189, 315, 322, 388), and
the foam, although complex, is essentially pro-
teinaceous (247).
Foaming of the digesta is common: normal-
ly, however, such foam is of low persistence
and is in only small amounts. In bloat the foam
is rigid and of high persistence (52, 206, 208,
247) and collects in large amounts. That the
ruminant has difficulty in coping with exces-
sive foaming of its stomach contents is support-
ed by the association of foaming and feedlot
bloat and by the experimental production of
bloat when foaming of the digesta is induced
by administration of egg white (44).
In contrast to free-gas bloat, where the sep-
aration stage of eructation appears normal, in
foamy bloat persistent foam indicates a marked
prolongation of the separation stage. Transfer
]OU~,~AL O F DAII:Y SCIENCE VOL. 57. :No. 7
of the free gas to the esophagus also is im-
paired in foamy bloat because the effective in-
crease in the digesta volume increases the diffi-
culty of clearing the cardia. This situation is
aggravated by the apparent inability of the
animal to expel foam: foamy digesta entering
the esophagus induces reflex swallowing and
only reaches the mouth in minor amounts
which are promptly re-swallowed.
Three general requirements for the develop-
ment of persistent foams in the ruminal digesta
can be recognized. (1) The balance of surface-
active materials must favor foaming. (2) The
environment provided by the digesta must fa-
vor foam persistence; that is, factors such as
pH, temperature, ionic composition and
strength must lie within the optimal ranges.
(3) There must be an adequate, but not neces-
sarily excessive, gas production. These require-
ments are apparently satisfied partly by the
plant, partly by the animal, and partly by the
activities of the rumen microorganisms. The
foam is, in effect, the result of a complex inter-
action of plant, animal, and microbial factors.
Components of the foaming system are logi-
cally sought in the foam itself. Until recently,
this direct approach seems to have been tried
on few occasions (26, 168), presumably be-
cause of the experimental problems. Bloat
foams typically are heavily contaminated with
rumen liquor, plant debris, and microorgan-
isms, making detection of small but possibly
critical differences in composition, structure,
or distribution difficult. Further, the foaming
system is readily altered or even destroyed dur-
ing handling. Since the initial work of Mangan
(246, 247) there have been many studies on
the generation and properties of bloat and
bloat-like foams, utilizing a wide variety of
methods (33, 76, 150, 166, 176, 184, 185, 219,
220, 221, 239, 313, 340, 381, 397, 411).
The first biophysical consideration of foams
in the rumen was by Nichols (288). Recently,
physical aspects of protein foam formation and
bloat foams have been investigated by Laby
(229, 230), Jones (206, 208, 210, 211, 213),
and Buckingham (52, 53). Although the
pbysico-chemical requirements for stable foam
formation cannot be dissociated from any
study of agents contributing to bloat, further
discussion of the actual physics of foam forma-
tion is outside the scope of this review.
The contributions of the plant, animal, and
microbial factors to the bloat foam are dis-
cussed below.
The Contribution of the Plant
Many plant materials, apart from protein,
 R E V I E W : FOAMY BLOAT
have been suggested as contributing to the oc-
currence of bloat foam. Most favored have
been saponins (235), pectic substances (95),
and pectin methyl-esterase (156) while hemi-
celluloses (95, 168) and nonvolatile acids
(294, 338, 339) also have been suggested as
important. Chloroplast lipids (247), galacto-
lipids (13), and sulfolipids (341) have been
investigated with regard to possible antifoam-
ing activity. In general, however, substantive
correlations between chemical composition and
the ability of the plant to cause bloat have
proved elusive.
Care is needed in the interpretation of
chemical analyses of bloat-provoking pastures.
Many plant constituents increase with lushness
and maturity to a certain stage and then de-
cline as the plant ages further (56). Because
of the general correlation between bloat and
lushness, there is also a correlation -- some-
times high - between bloat and such constitu-
ents (25, 266). It does not follow, however,
that a high correlation between bloat and a
partictdar constituent necessarily indicates sig-
nificant involvement of that constituent in the
occurrence of bloat, a point overlooked by
some workers.
Explanations for the variable nature of bloat
have been sought in fluctuations in many plant
components (81, 266, 273). However, while
such fluctuations may explain long-term
changes in the occurrence or severity of bloat,
they seem unlikely to explain the short-term
variation which is a characteristic feature of
the disorder. Although there are reports that
both cattle and sheep bloat more severely in
the afternoon than in the morning (103, 315,
363), this is certainly not an invariable occur-
rence (189).
It is not yet possible to predict whether a
particular pasture will provoke bloat.
Minerals. Fertilizer treatment and soil min-
eral content have been claimed, in many pa-
pers, to affect bloat. The data, however, are
confusing and often contradictory. In many in-
stances the full consequences of fertilizer treat-
ment in plant and animal response do not ap-
pear to have been appreciated. While fertilizer
requirements will vary from district to district
and from farm to farm, as a generalization, any
fertilization program that increases the vigor
of legumes or legume content of pasture will
increase incidence of bloat, ,and decrease en-
courages decrease. High rates of application
of phosphate, for example, encourage grass
dominance and reduce bloat (345, 377, 404).
High potash fertilization of alfalfa has reduced
bloat incidence (55, 257).
757
Legumes contain higher Ca and Mg than
grasses (373), and frothy rumen contents con-
tain more magnesium ammonium phosphate
than nonfrothy ingesta (302).
Smith and Woods (354, 355) showed that
spraying with Ca and Mg salts increased sever-
ity of bloat and that intraruminal administra-
tion of chelating agents reduced severity. The
chelating agent, ethylenediamine di-orthohy-
droxyphenyl acetic acid, was bacteriostatic and
depressed both forage intake and ruminal pro-
duction of volatile fatty acids (362). Warner
et al. (392), by spraying urea and calcium car-
bonate on alfalfa, raised intakes by lambs of
both N and Ca and found that both increased
incidence of bloat.
Studies of mineral content of legumes have
been by many workers, but no correlations
with bloat have been consistent (96, 181, 257,
273, 276). Stifel et al. (364, 365) in a study
of alfalfa leaf chloroplasts and bloat showed
a positive correlation of bloat with leaf Mg and
a negative correlation with leaf Ca. In a more
recent study, Miltimore et al. (275) demon-
strated a negative correlation with Mg but a
positive one with Ca, Ni, and Zn. The latter
correlation supports the observation of Harris
and Sebba (166) that there is more Ni and
Zn in strong than in weak alfalfa foams. Cal-
cium has assisted the development of strong
protein and saponin foams (206, 247) ,and is
involved in the formation of pectates in the ru-
men (409). It may be relevant that a high
Ca/P ratio has favored the occurrence of bloat
(98, 99, 100).
Saponim. The place of saponins in bloat has
been controversial. Two roles have been pro-
posed, pharmacological and surface-active:
both have been discussed in previous reviews
(81, 82, 190, 191). Cheeke (64) recently re-
viewed the pharmacological, nutritional, and
physiological implications of saponins. Hun-
gate (175) lists many of the compounds, other
than saponins, that have been considered phar-
macologically important in bloat. The general
conclusions are that it is not possible to corre-
late bloat with respiratory inhibitors or any
other toxic components of plants (123, 175,
181, 182, 258).
The possibility that saponins were a major
component of foams in foamy bloat seemed un-
likely following the foam studies of Mangan
(247). Since then it has been accepted that
saponins do not play a major part in the
genesis of bloat foams (175, 250) although
there is still some interest in their effect (39,
150, 177, 300). It may be significant that some
rumen bacteria are capable of rapidly degrad-
JOURNAL OF DMRY SCIENCE VOL. 57, NO. 7
758 CLARKE AND R E I D
ing dietary saponins (157, 158, 159).
Pectins. Conrad and colleagues (95) and
Head (168) first proposed pectins as being
capable of causing the stable foam in pasture
bloat. According to Conrad et al. (95), pro-
duction of the foam was associated with a high
rate of gas production from the pectin-contain-
ing fibrous fraction of the feed. However, Boda
and Johns (42) showed that the rapid gas pro-
duction which follows the ingestion of legumes
results from fermentation of plant juice rather
than fiber.
Pectic substances may constitute up to 8%
dry weight of some forages (95, 231, 312, 386,
409). Marked daily variation of the amounts
in alfalfa, and a positive correlation of the
amounts with bloat, was found by Pounden
and Conrad (310), but Pressey et al. (312)
could not correlate bloat with either water-
soluble or total pectins although added pectin
increased foam stability of alfalfa extracts in
vitro (313). Pectin is degraded rapidly by ex-
tracellular enzymes and microorganisms and
does not accumulate in the rumen (106, 313,
401, 408, 409).
Increased viscosity, of the digesta in legume
bloat was ascribed to the activity of pectin
methyl-esterase by Gupta and Nichols (156,
289, 291): the enzyme is in aqueous extracts
of fresh alfalfa, in rumen fluid, and, with poly-
galactnronase, in rumen bacteria and protozoa
(401, 408, 409). Demethylation of pectin by
the esterase greatly increased the viscosity of
rumen contents incubated in vitro, and it was
posttilated that the action of the enzyme in
vivo assisted pectic gel formation and entrap-
ment of gas. Correlation between bloat inci-
dence and intake of the enzyme by the dnimal
has been positive (292, 293): enzyme activity
in the plant was higher although the dry
weight of the plant was lower on days when
bloat occurred. It was also demonstrated
(290.) that bloat in cows feeding on legumes
could be inhibited by feeding an inhibitor of
pectin methyl-esterase, alkyt aryl sulfonate.
The protection lasted for approximately 24 h,
and it was suggested that the material would
be suitable for use in capsules designed for
controlled release of the agent in the tureen.
Alkyl aryl sulfonates are common detergents:
both Barrentine et al. (19) and Reid and
Johns (330) found them unreliable in the
treatment of bloat and having no long protec-
tive effect. A naturally occurring inhibitor of
pectin methyl esterase, found in rumen con-
tents by Penn, Deese and Nichols (301), was
suggested as a contributing factor in prevent-
ing bloat in nonbloating animals.
JOURNAL OF DAIRY SCIENCE VOL. 57, NO. 7
Although pectins are not primary foaming
agents, the current theory is that they increase
viscosity of rumen fluid and could act as foam-
stabilizing agents in bloat (206, 409). Jones
(206) recently showed that sodium pectate
will stabilize the foam produced by leaf pro-
tein; however, Wright (409) suggested that
pectate formation might reduce the amount of
Ca available for stabilizing protein foams.
Proteins. The first major evidence that plant
proteins were the p~nary foaming agents in
bloat came from a study of the pH optima for
maximum foam strength (247). Solutions of
saponins exhibited maximum foam strength at
pH 4.4 to 5.5 whereas the maxima of solutions
of cytoplasmic protein were in the range of pH
5.5 to 6.0 depending on salt concentration.
Foams generated from rumen fluids of clover-
fed cows had maximum foam strength between
pH 5.4 and 5.7. At the onset of clover bloat
rumen fluid may have a pH as high as 6.6, but
in severe bloat it usually is in the range pH 5.6
to 5.9.
The major role for protein was supported by
Bartley and Bassette (26), but Head (168)
concluded that the foaming constituent was
largely carbohydrate. Foaming properties of
red clover cytoplasmic proteins have been
similar to those of proteins of alfalfa (150).
The soluble leaf proteins of higher plants
may be divided into two main groups, Fraction
1 and Fraction 2, on the basis of molecular size
(351). Fraction 1 is a homogeneous protein
with a molecular weight of approximately
550,000 and has been identified as the enzyme
ribulose diphosphate carboxylase (249, 378).
It is the major protein in the chloroplasts of
green leaves (244). Fraction 2 is a mixture of
proteins of molecular weight 10,000 to 200,000
and includes all soluble leaf proteins other
than Fraction 1 (351).
In 1962 McArthur (251) stated that the
basic cause of foam formation in legume bloat
is the presence in the rumen of a surface-de-
natured cytoplasmic protein of legumes. This
protein was later, isolated from alfalfa protein
extracts as a homogeneous protein with a sedi-
mentation coefficient similar to 18S or Fraction
1 protein (255).
McArthur, Miltimore, and Pratt (255) listed
the events which, they suggested, probably
lead to foam formation with Fraction 1 pro-
tein. Following release of chloroplasts in the
rumen, Fraction 1 protein rapidly enters the
rumen fluid, the protein in solution as an al-
most spherical particle. If the molecule reaches
the surface without undergoing microbial de-
gradation, it uncoils there and becomes insohi-
 R E V I E W : FOAMY B L O A T 759

ble. It is this surface-denatured protein that to misinterpretation.

stabilizes the foam. Foam stability is reduced It has been suggested that differences in re-
by agitation as the protein then coagulates be- suits with Fraction 1 from various legumes
cause of the strong cohesive forces between have arisen because of the different properties
molecqfles. The observations on denaturation of the various isolated fractions. However,
reported by McArthur et al. (255) were sup- Jones and Lyttleton (212) have stated that
ported by the findings of Jones and Lyttleton Fraction 1 has similar properties whatever its
(208). Stifel et al. (364) found a high correla- origin: traces of phenolic compounds adhering
tion with bloat of Ca and Mg binding to Frac- to the protein cause the different electro-
tion 1 chloroplast protein and, following the phoretic mobilities with some preparations.
observations of McArthur et al. (255), sug- In a study of leaf proteins of both red and
gested that any factors which promoted pro- white clover, Jones and Lyttleton Showed that
tein denaturation would enhance bloat. The Fraction 2 proteins, as well as Fraction 1, wero
metal-binding they reported could salt-out surface-denatured by foaming at pH 5.8 and
Fraction 1 protein thereby stabilizing the that, as in the case of Fraction 1, Fraction 2
rttmen foam. Surveys in Canada had positively proteins also produced strong foams over the
correlated nitrogen and protein in forage with range of pH which exists in the rumen when
bloat incidence (273, 276) although authors bloat occurs (78, 208, 210, 253). The total
admitted the conclusions in the latter work soluble leaf proteins of red and white clovers
were of doubtful value because the samples produce foams of maximum strength at pH 5.4
were all of alfalfa at the hay stage of develop- to 6.0 (210, 229, 247). Both Fraction 1 and
ment. In a survey of alfalfa varieties, McArthur Fraction 2 proteins have their maxima for foam
and Miltimore (252) showed that Fraction 1 strength within this pH range, Fraction 1 hav-
protein comprises 3.9 to 5.3% dry weight of ing the sharper maximum at about pH 5.8
red clover and alfalfa and that the content of (210). Both protein fractions contribute to
this protein increased with age to a maximum properties of foams in legume bloat (213).
before decreasing to a constant amount. Trefoil Tannins. About the time that Cooper,
(Lotus corniculatus), a nonbloating forage, Eslick, and McDonald (97) showed that the
contained about .5% Fraction 1. In a later pa- bloat potential of forages was positively corre-
per (254), t~ve bloating forages contained lated with the volume of foam that could be
more Fraction 1 and total protein than six non- generated from extracts of leaf, Kendall (220)
bloating forages. The contents of Fraction 1, showed that foam production in vitro was in-
lipoprotein, and total protein in trefoil, white hibited by tannins. The ability to foam was re-
clover, and four varieties of alfalfa were exam- stored by addition of polyvinylpyrrolidone
ined by Rommann et al. (337). Minor differ- (PVP), an agent known to complex with tan-
ences in Fraction 1 protein were not sufficient- nins and polyphenolics (7). This agent also in-
ly large to be a primary factor in the etiology creased the volume of foam obtainable from
of bloat. nonbloating forages. Conversely, the addition
That Fraction 1 was important in alfalfa of a commercial tannin to the extract of a
bloat was supported by Stifel et al. (365), who bloating legume reduced the foam. Kendall
showed that total and soluble leaf chloroplast postulated that with nonbloating legumes,
protein was directly related to the severity of foam production in the rumen might be limited
bloat in sheep and cattle, with the highest cor- by tannins in the plant.
relation between Fraction 1 chloroplast protein Bloat has been reported as occurring on only
and bloat. The threshold of Fraction 1 protein one of the tropical legumes in Australia (164),
for incidence of alfalfa bloat in Canada was re~ but this absence of bloat could not be corre-
ported to be 1.8% by Miltimore et al. (275). lated with the tannin content of the leaves ac-
Doubt has been cast on the importance and cording to Hutton and Coote (178), who
absolute contribution of Fraction 1 to the foam examined 19 species of legume.
in bloat by Jones and Lyttleton, who criticized Jones, Lyttlcton, and Clarke (214) showed
the conclusions of Stifcl et al. (364, 365) on that of four forage legumes, the three that
the grounds that the methods yielded only neg- cause bloat, namely red, white, and subter-
ligible amounts of Fraction 1 (208). They also ranean clovers, all released high soluble pro-
pointed out that the original conclusions of tein when extracted with buffer. The fourth
MeArthur, Miltimore, and Pratt (255) on the legume, lotus major (Lotus pedunculatus),
importance of Fraction 1 were based mainly which does not cause bloat, released large
on the analysis of leaf proteins by gel filtration, quantities of soluble protein only when PVP
a method which can be deceptive and liable was in the extracting buffer. Jones and Lyttle-
1OURNAL OF DAIRY SCIENCE VOL. 57, NO. 7
760 CLARKE AND REID
ton (209) in a recent study of 21 temperate
and tropical legumes demonstrated a strong
positive correlation between absence of protein
precipitants and tendency to cause bloat, a cor-
relation that was also demonstrated by Jeffers
et al. (188). Cope, Bell, and Smart (101) re-
ported a leucoanthocyanin polymer from Les-
pedeza that could inhibit some plant enzymes
and suggested that pectin methyl-esterase
might be inhibited in vivo with a consequent
decrease in bloat. Cheeke (64) suggested that
tannin-saponin complexes may partly explain
the low-bloat potential of high-tannin legumes.
Confusion has arisen over use of the word
tannin. Tannins are polyphenolic compounds,
with a molecular weight of 500 to 3,000
(371), which can combine with and precipi-
tate proteins. The most efficient for this pur-
pose, condensed tannins, have recently been
identified as polymeric flavanols, also known
as flavolans (207). They appear to be unspe-
cific in their action. Those of Lotus peduncula-
tus precipitate leaf protein, rumen protozoal
proteins, salivary mucoprotein, and the foam-
ing agents in rumen fluid from bloating ani-
mals (206).
According to Jones (206), most measure-
ments of tannins in forages have been of total
phenolic compounds or of leucoanthocyanins
which do not precipitate soluble proteins. Red
clover, in common with most pasture legumes,
has a high content of polyphenols which do
not precipitate the leaf proteins. Jones and
Lyttleton (209) suggested that the positive
correlation between Fraction 1 protein and the
tannin content of alfalfa found by Miltimore
and colleagues (275) resulted from the latter's
assay method which could not distinguish be-
tween condensed tannins and leucoanthocya-
nins. Jones (206) could not find condensed
tannins in alfalfa.
The breeding of tannins into existing le-
gumes or the establishment of agronomicafly
suitable tannin-containing legumes in mixed
pastures with clovers and alfalfa might be
profitable approaches to bloat prevention
(206, 207, 214, 299). A recent examination
of 44 Trifolium species for plants that could
be used in a plant breeding program demon-
strated condensed tannins in only T. arvense
and T. affine (207).
Lipids. IApids, particularly those of chloro-
plasts, were suggested by Mangan (249) as
being important antifoaming materials in the
rumen. Removal of chloroplasts, which contain
glycolipids (36, 394, 396), increased the sta-
bility of tureen foam in vitro; removal of lipid
from the chloroplasts also increased stability
JOURNAL o r DAIRY SCIENCE VOL. 57, N O . 7
of the foam. However, chloroplasts may have
a dual role in bloat - as particles stabilizing
the foam and as carriers of antffoaming agent
(247). Oxford (298) showed that the ciliate
Epidinium could aid foam formation by rapid
removal of whole chloroplasts from the rumen
liquid.
Glycolipids constitute a significant propor-
tion of the lipid fraction of red clover (395).
Bailey (13) investigated lipid-bound sugars
of red clover but found no correlation of these
constituents with bloat. Galactolipids were
readily released into tureen liquid and were
rapidly degraded (13), both by bacteria and
protozoa (14). There appeared to be variation
from animals in the rate of release, but this al-
so could not be correlated with bloat (13).
Bryant (49) investigated release of galacto-
lipids from the plant by chewing. Russell and
Bailey (341) studied both lipid-bound galac-
rose and sulfoquinovose in red clover. The sul-
folipid was highly surface-active, but the au-
thors considered the concentration in the ru-
men unlikely to reach where foam was inhib-
ited, even though sulfolipid was hydrolyzed
at a slower rate than galactolipid. It is thought
that the action of penicillin in preventing bloat
may result partly from inhibition of bacteria
that modify lipid (248, 410).
Fraser (146) could not find any variation
in the lipid content of ryegrass, red and white
clover which would correlate with bloat al-
though there was always higher lipid in grass
than in clover. No differences were significant
in amounts of various lipid fractions in rumens
of bloating and nonbloating clover-fed cows
(146).
Lipids are in appreciable amounts in mmen
contents of legume-fed cows, but the exact na-
ture of their association with bloat remains to
be elucidated. Wallace (390) found 11 to 24%
lipid in rumen foam from bloating an'hnaals
feeding on fresh alfalfa. Yellow bubbles in the
froth were coated with and had a high content
of particulate matter in their surface layer.
Laby and Weenink (230) used foam separa-
tion to examine the surface-active components
of rumen liquid taken before feeds of bloating
and nonbloating red clover. Differences in non-
acid lipid and fatty acids were in foam but
were not refected in tureen liquors. The yel-
low bubbles previously described by Wallace
(390) were separated and contained more
than 30% lipid. They were in greater concen-
tration in cows about to bloat than in cows that
did not bloat at the following feeding.
A significant contribution was made by Sti-
fel, Vetter, and Mien (364), who proposed a
 R E V I E W : FOAMY BLOAT
competitive relationship between lipid and
protein in alfalfa foams. According to these
authors, polar lipids compete for metal ions
which bind to Fraction 1 protein, thus decreas-
ing protein denaturation and reducing foam
formation. It was also suggested that when the
saturated fatty acid content of the phospho-
lipids increases, as occurs during maturation
of alfalfa (224), calcium binding increases at
the expense of Fraction 1 and bloat is reduced.
Conversely, with immature alfalfa, with phos-
pholipid high in unsaturated acids, more cal-
cium is available for binding to protein, in-
creasing denaturation, surface viscosity, and
bloat. In a seeond paper Stifel et al. (365)
showed there was a strong negative correlation
between alfalfa chloroplast lipid and bloat.
Miltimore and colleagues found bloat was posi-
tively correlated with the lipids of alfalfa
(275).
From studies of in vitro foaming systems,
together with an examination of tureen fnams
after freeze-etching, Buckingham (53) has
suggested that apart from, and seemingly in
opposition to, their role as antifoaming agents,
lipids may have an important function in pro-
moting aggregation of protein molecules in
rumen foams, thus increasing foam stability.
Jones (206) examined the polar lipids of red
clover leaves for their effectiveness as anti-
foaming agents on Fraction 1 protein foams
generated in vivo. Antifoaming potential de-
creased in the order phosphatydyl choline, di-
galactosyl diglyceride, monogalactosyl digly-
eeride, sulfolipid. The effect of these plant lip-
ids was to reduce the persistence and rigidity
of the protein foams. With protein and lipid
as foaming and antifoarning agents respective-
ly, efficient foam-stabilizing agents also would
be needed to explain the highly persistent na-
ture of rumen foams.
Another role for lipids was proposed by Ko-
marek (227), who suggested that their benefi-
cial effect in preventing bloat resulted from
their ability to form complexes with toxic com-
pounds responsible for bloat. In view of the
success of antifoaming agents of widely differ-
ing chemical nature in both treating and pre-
venting bloat, this hypothesis seems untenable.
The Contributions of the Animal
Animal factors in bloat have been discussed
in earlier reviews (78, 81, 82, 190, 191), by
Johns, Mangan, and Reid (194), and by Men-
del and Boda (260). Here, only selected topics
will be discussed, with particular emphasis on
individual susceptibility and on the contribu-
tion of saliva.
761
Susceptibility. There is great variation in
susceptibility of animals to bloating feeds,
ranging from animals which bloat readily and
severely to those which bloat seldom ff at all
(17, 78, 165, 176, 189, 225, 236, 260, 261,
322, 328). Clarke and Reid (78) used the
terms HS and LS to denote animals at either
extreme of the range, i.e., animals of high or
low susceptibility. If an animal of low suscepti-
bility does bloat, its rumen contents are foamy
and appear indistinguishable from those of a
bloating animal of high susceptibility (322).
It is important that the susceptibility of all ex-
perimental animals be established (78, 260);
bloat-limiting factors may not be the same in
animals of different susceptibility when they
are not bloating (78). Many HS animals die
of bloat before reaching maturity (80); those
that survive may well have adapted their be-
havior to avoid the worst risks, Animals that
have given little or no previous indication of
being susceptible may bloat severely and die
when they first suffer from the disorder. Wide-
spread use of bloat-preventing agents has
made recognition of susceptibility more diffi-
cult.
Animals appear to have an intrinsic suscep-
tibility to bloat that is determined genetically.
Evidence indicating a genetic basis includes
(1) the prevalence of bloating in progeny of
particular sires (165, 189, 191, 194, 225, 328),
(2) the similarity in the bloating behavior of
monozygotic twin sets (165, 189), and (3)
differences in susceptibility of cattle of differ-
ent breeds (205, 270, 295, 328). Differences
between breeds are relative rather than ab-
solute. It is often claimed that tropical breed~.
of cattle are less susceptible to bloat than are
temperate breeds and that some (e.g., Santa
Gertrudis cattle) are even immune to the dis-
order (205, 327); nevertheless, losses by death
from bloat occur in tropical breeds, including
Santa Gertrudis (I. Saxton, personal eommurti-
cation).
The immediate site of action of the factor
or factors determining susceptibility lies within
the contents of the rumen. In animals with a
rumen fistula, total exchange of these contents
between an HS and an LS animal before feed-
ing, results in a temporary exchange of sus-
eeptibilities (78). The effect lasts some 24 h.
before the host animal reimposes its innate sus-
ceptibility.
The identity of the susceptibility factor or-
factors is obseure. Mendel and Boda (260) list
the following as possible animal factors: "sail-
vary flow, salivary composition, fluid transfer
from the rumen, rttmen motility (particularly
IOURNAL OF DAIRY SCIENCE VOL. 57, NO. 7
762 CLAnKE AND aEtD
with respect to eructation efficiency), gas pro-
duction, microbial population, rate of eating,
rumen pH, and breakdown products of nitro-
gen metabolism (especially soluble protein)."
Other factors which could be added to that list
are the pattern of transfer of particulate matter
out of the rnmen, the "set" (effective sensitiv-
ity) of the sensory nerve-endings of the gut
wall, the permeability of the rumen lining
epithelium, endocrine activity (including its
effects on the microbial population), and the
effectiveness of acid-base regulation. In a study
of differences between susceptible and nonsus-
ceptible cows fed fresh alfalfa, Mendel and
Boda (260) found significant differences only
in (a) salivation during rest and feeding, sus-
ceptible animals secreting less saliva but with
a higher bicarbonate content, and (b) the
physical nature of rumen contents. They could
find no differences in salivary Na, K, PO4, and
mucin; frequency and amplitude of rurnen
contractions; rate of gas production; total, solu-
ble and ammonia nitrogen of rumen liquid;
buffering capacity of digesta and saliva; rumen
pH and water consumption. In a similar study
in New Zealand, still in progress, in whidh
comparisons have been of HS and LS cows fed
fresh red clover or clover or lucerne hay, no
consistent differences have been found so far
in salivation rate during feeding, water flows
through the rumen, dry weight of rumen con-
tents before feeding, their buffering capacity
before and after feeding, microbial activity in
them before, during, and after feeding, and
rumination behavior (328).
According to Johns (189) and Hancock
(165), the rate of eating is not important. The
relative rate of release of foaming and anti-
foaming agents into the rumen from chewed
feed was suggested as a factor in bloat (321,
332); an apparent animal difference in effec-
tiveness of chewing with regard to protein re-
lease has been found (49), but this has not
been related to susceptibility. The effects of
exchanging contents of the rumen (78), re-
ferred to above, also suggest chewing and sali-
vation are not important in the short term.
That conclusion is supported by effects of ex-
changing boll of chewed and ensalivated feed
rather than rumen contents. When boll were
collected by hand as they were delivered
through the cardia of an LS animal and then
inserted into the tureen contents of a fasting
HS animal, the HS animal bloated, but when
US boll were transferred to LS recipients, the
LS animals did not bloat (78).
Knowledge of the origins of differences in
susceptibility could provide a rational basis for
JOUI~NAL OF DAIRy SCIENCE VOL. 57, NO. 7
the breeding of cattle with a reduced suscepti-
bility to bloat (328). Establishment of such
low bloating strains seems a practical goal. In
dairy cattle, bloat susceptibility appears to be
dissociable from productivity (328), suggest-
ing that it should be possible to reduce sus-
ceptibility and at the same time retain high
productivity. If causes of low susceptibility
could be established or a marker that corre-
lates with low susceptibility be found, the need
to determine susceptibility by feeding trials
might be avoided. A considerable shortening
of breeding programs would result.
Saliva. A copious salivary secretion is one
of the characteristic features of the rtmainant
digestive system, the dailv seeretion in cattle
being variously estimated" as 25 to 190 liters
(11, 66, 175, 217, 267, 413) and in sheep as
1 to 24 liters (66, 217).
Saliva is secreted by a number of glands
(54, 216, 217), and its composition varies ac-
cording to its origin. Basically, the glands and
their secretions may be classified as serous,
mucous, and mixed (216, 217). Serous saliva
is thin, high in bicarbonate, and strongly buf-
fered but is low in protein and contains very
little mucoprotein. It is secreted continuously
although at varying rates whether the animal
is feeding, ruminating, or resting. Mucous
saliva is thicker, generally containing consider-
able protein and mucin. It is secreted predomi-
nantly during feeding, little usually being se-
creted at other times. Salivation is under re-
flex nervous and hormonal control (9, 40, 54,
63, 68, 94, 215, 216, 217, 218, 343). The na-
ture of the feed itself influences the amounts
and composition of the saliva secreted during
feeding (9, 11, 12, 43, 128, 217, 297, 397,
413). Other important influences on salivation
are the reflex effects arising from stimulation
provided by the digesta in the gut (68, 94,
217). Yet other influences are the mineral sta-
tus of the animal, especially its Na status (40),
and the osmolality of the extracellular fluid
(63).
Aspects of salivation likely involved in bloat
include: volume secreted, buffering power,
amount of bicarbonate secreted, and amounts
of mucoprotein and mucin secreted.
Cole and Mead (85, 86, 259), 30 yr ago,
proposed bloat was due to a "physical deficien-
cy of the diet." They considered that diets
causing bloat contained insufficient roughage
to stimulate eructation. Following his studies
of eructation (398) and the discovery that me-
chanical stimulation of the rumen walls could
cause salivation as a reflex response (68),
Weiss also proposed a physical deficiency lay-
 REVIEW:
pothesis based on the premise that bloat was
a consequence of reduced salivary secretion
(397). In experiments with sheep fed freshly
cut lucerne (397), he found bloating was asso-
ciated with increased viscosity of the tureen
contents. He also found that if chaffed grass
hay (10% by wet weight) was mixed into the
lucerne, bloat did not occur, and that introduc-
ing chaffed hay through a rumen cannula re-
duced the viscosity of the tureen contents of
sheep fasted overnight. He attributed the ef-
fects of chaff to reflex stimulation of salivary
secretion. From these experiments, Weiss
postulated that succulent lucerne did not pro-
vide a strong stimulus for salivation so that less
saliva was secreted, resulting in an increase in
the viscosity of the contents of the rumen, and
foaming.
That cattle salivate less when they eat suc-
culent young legumes than when they eat
more mature plants has been confllTned by
Meyer et al. (267), who also found an inverse
relationship between salivation and water con-
tent of the feed; results have been similar at
this laboratory.
Effects of manipulation of dietary roughage
appear to support the physical deficiency hy-
pothesis, whatever the mechanisms involved.
In general, decreasing dietary roughage in-
creases the risk of bloat; grinding increases the
incidence of bloat on dry diets (84, 91, 171,
259, 314) while at this laboratory mincing the
feed has induced bloating in LS cows fed
fresh red clover. Conversely, increasing dietary
roughage - by feeding mature rather than im-
mature legumes, using mixed grass/legume
pastures, by adding chaffed or long hay to the
diet - decreases the risk of bloat (86, 89, 92,
397). In both situations, however, there are
effects other than any there may be on saliva
secretion, and which may weft be involved in
the responses. Thus, while finely (but not
coarsely) ground diets reduce salivation (217,
402), they are also associated with reduced
rnmen motility (91, 92, 324), increased gas
formation (171), and probably an increase in
particulate matter capable of stabilizing foams.
Again, although hay feeding results in a free
flow of saliva (12, 217, 267), it also tends
to strengthen rumen motility (91, 92, 323,
324), and can improve eructation efficiency
(90). Feeding hay supplements has not
proved a reliable preventive measure - to be
effective, large amounts of scabrous hay must
be eaten (86, 89, 108, 169, 189).
Mendel and Boda (260) reported that
bloat-susceptible cattle secrete less saliva dur-
ing resting and feeding than nonsusceptible
FOAMY BLOAT 768
cattle, but Meyer et al. (267) were unable to,
demonstrate this because of large day-to-day
variations, attributed to variations in water
content of the feed.
Distention of the rumen during bloating
may stimulate salivation initially, but con-
tinued distention is likely to be inhibitory (9,
217, 218, 307); on the other hand, distention
of the esophagus is likely to stimulate secre-
tion (217, 218, 307), especially of mucous
saliva. Phillipson and Mangan (306) reported
a fivefold increase in the mueoprotein con-
tent of submaxillary saliva of anaesthetized
calves following inflation of the rumen.
Another factor that could affect salivation is
the change in physical nature of the rumen
contents during bloating.
Saliva contributes bicarbonate and phos-
phate which help buffer pH changes in the
rumen. Additional buffering action occurs by
the absorption of volatile fatty acids through
the rumen epithelium and simultaneous secre-
tion of bicarbonate into the rumen (8, 358).
No differences have been found in buffering
capacity of the contents of HS and I_~ cattle
before or after feeding (260, 328). Mendel
and Boda (260) found bloat-susceptible cows
secreted saliva with a higher bicarbonate con-
tent (although less in volume) than nonsus-
ceptible cows.
Acidification of bicarbonate is an important
source of gas in the rumen. A study of acid-
releasable COL in ruminal fluid has been made
by Gupta et al. (155). The daily production
of CO2 from bicarbonate in saliva and enter-
ing the tureen during VFA absorption
may be greater than 250 liters. This figure is
not particularly relevant to bloat even when
fermentation gas is added; the important meas-
ure is the peak gas production at the time of
maximum foam strength.
In the past 16 yr there has been increasing
interest in the role of saliva (20, 33, 78, 189,
206, 242, 243, 247, 322), and it is now known
to exhibit both foaming and antffoaming
properties. This is in line with the fact that
compounds that are foaming agents in one
situation, or by themselves, can be antffoam-
ing agents in another situation.
Working with bovine saliva collected from
under the tongue by suction during feeding,
Lyttleton (242, 243) showed that 41 to 66%
of the protein contained sialie acid (N-aeetyI
neuraminic acid). This mucoprotein, of which
sialie acid constitutes 20 to 30% (153, 296)
is responsible for the thick nature of mucous
saliva, the strong negative charge of the sialie
acid-containing side-chains imparting a high
JOURNAL OF DAIRY SCIENC/~ VOL. 57, N O . 7
764 CLARKE A N D R E I D
degree of viscosity to the protein (152). Jones
(206, 213) has found a desialated mucopro-
tein of saliva is a major component of the solu-
ble foaming constituents. The protein content
of the saliva collected by Lyttleton (242)
ranged from .9 to 2.2 (mean ----- 1.69) mg/ml.
Clarke, Hungate, and Reid (unpublished re-
suits) collected saliva from an esophageal
t~stula or from the cardia, in both cases during
stimulation by distention of the esophagus with
a balloon. This saliva had marked foam-
stabilizing properties when incubated with
rumen liquid and minced clover in vitro (76,
78). It was more viscous and ropier in nature
than that collected by Lyttleton (242), yet
protein was considerably lower (.2 to .9; mean
.5 mg/ml). Jones (206) showed this type of
saliva was low in mucoprotein and contained
a carbohydrate of high molecular weight as
well as a number of proteins of low molecular
weight. In addition, he verified its foam-
stabilizing activity with Fraction I protein
foams (206). Lippke (237) also found that
salivary mucins increased foam stability. Men-
del and Boda (260) could not demonstrate
differences in the mucin content of saliva from
HS and LS animals.
In contrast to the above foam-stabilizing
properties of saliva, Bartley and co-workers
demonstrated in saliva an antffoaming agent
which they identified as salivary mucin. Van
Horn and Bartley (381) showed that addition
of saliva to frothing rumen contents in vitro
released gas from the foam, and Bartley and
Yadava (33) showed that the antifoaming ac-
tivity was also effective on saponin foams. Fol-
lowing this it was postulated that bloat is in-
creased by namen mucinolytic activity (137),
and rumen contents of HS animals exhibited
higher mucinolytic activity than those of LS
animals, both aerobic (282) and anaerobic
(137, 281) rumen bacteria being responsible.
The theory was proposed, therefore, that saliva
is an anti'foaming agent and that animals sus-
ceptible to bloat secrete less saliva than LS
animals (260) and have more active popula-
tions of mucinolytic organisms which effective-
ly destroy the antifoaming properties of the
saliva (137, 281). However, there may be an
additional explanation for these observations.
If desialated salivary mucoprotein is a foam-
stabilizing agent, as proposed by Jones (213),
addition of mucinolytic bacteria to the tureen
could accelerate the production of foam-stabi-
lizing material from whole mucoprotein as well
as destroy antffoaming activity. The full role
of saliva in bloat has yet to be determined;
the fates of salivary mucin and mucoprotein
~JOURNAL OF DAIRY SCIENCI~ VOL. 57, NO. 7
certainly warrant further investigation. The
relative proportions of the many protein com-
ponents of saliva have shown some promise as
a marker for LS cattle (Clarke, R. T. J., W. T.
Jones, and C. S. W. Reid, unpublished re-
suits).
The digesta. When HS cattle are kept on a
bloat-provoking diet, their rumen contents are
seldom free from foam. Even after 24 h of fast-
ing, a myriad of small bubbles is typical. "Hie
bubbles give HS rumen contents a lighter color
and the appearance of greater digestion as
compared with LS contents (175, 356).
Bloat-susceptible cows, in the study of Men-
del and Boda (260), had a higher content of
dry matter in their ruminal digesta than did
nonsusceptible animals. Bartley (21) noted
that nonbloater twins had watery ruminal
digesta compared with those of bloater twins.
In recent work at this laboratory we have
found: (1) LS ruminal digesta after an 18-h
fast tend to contain a greater proportion of
coarse, fibrous feed residues; (2) HS digesta
contain a higher concentration of iqnely di-
vided particulate matter which could stabilize
foams (247, 389); (3) in contrast to observa-
tions of Mendel and Boda (260), the digesta
of HS animals after an 18-h fast usually have
a lower dry matter than those of LS animals
- this difference being partly but not wholly
related to feed intake (328). In other experi-
ments, a change in the nature of the rumen
contents of LS cows was induced by feeding
the animals for several days on minced fresh
red clover, which was eaten readily; coarse
fiber disappeared from the digesta and there
was a marked increase in the occurrence of
bloat.
Because of the foam in bloating digesta, the
specific gravity is lower, although the viscosity
is higher, than that of nonbloating digesta
(260, 288). Gutierrez et al. (162) showed
that ethanol-precipitable slime increased in the
tureen with onset of clover bloat. The slime
was composed of 61 to 64% protein, 8 to 14%
carbohydrate, and 7 to 10% ribonucleic acid.
Bartley and Bassette (26) gave a similar
anayslis for foam but found no nucleie acid.
Laby and Weenink, in analyses of prefeeding
rumen liquid by foam-separation (230), found
that the components of the foam may show
differences that are not retlected in the liquid
fractions. Foams from rumen liquid taken be-
fore bloating and nonbloating feeds of clover
showed differences in nonacid lipid, fatty acid,
and carbohydrate (230). Jones (206, 211)
has demonstrated that a major eomponent of
bloating foam is salivary mucoprotein and
 R E V I E W : FOAMY BLOAT
muein.
A net flux of water into the rumen across
the epithelium, leading to an accumulation of
water in the contents, has been proposed
(372) as a causative factor in bloat. No evi-
dence for such an effect has been found at this
laboratory.
Circulatory effects. Circulatory effects dur-
ing bloat and during experimental ruminal in-
sufflation have been studied by several workers
(34, 35, 104, 105, 284, 336). In general, the
observations can be accounted for by the circu-
latory and respiratory interference caused by
gastric distention, the development of hypoxic
hypoxia probably complicated by increased
CO2 absorption from the stomach, and distress
of the animal.
Changes in blood constituents. There have
been several studies of the effects of ruminal
insufflation on blood constituents (34, 116,
119, 370), but studies of effects of bloating are
fragmentary. Jackson et al. (182) showed that
cows grazing alfalfa pasture had greater
plasma cholesterol values than when they were
grazing blue grass and that the increase was
highly correlated with the severity of bloat.
Plasma ascorbic acid and red cell cholinester-
ase did not change with diet. Brown et al.
(47) found that I-IS animals had higher
plasma inorganic phosphorus. There was no
difference in blood ammonia, blood nonprotein
nitrogen, or hematocrit values between bloated
and nonbloated cattle. Hemoglobin and
methemoglobin levels apparently are not re-
lated to bloat (285); however, Beckett et al.
(34) observed that ruminal insufflation caused
increases in hemoglobin and packed cell vol-
ume in goats and calves. These workers also
found (34) raised K and suggested that hy-
perkalemia could be a contributing cause of
death in bloat.
Johnson, Dyer, and colleagues have studied
the metabolism of tryptophane, tyrosine, and
epinephrine in bloater and nonbloater cattle.
The bloater secretes less epinephrine (121,
202) but excretes more tyrosine (120) and
tryptophane (121, 202, 203) than the non-
bloater. Liver values for monamine oxidase
were less in bloaters (201). The data were
said to support the hypothesis of inborn differ-
enees in the metabolism of chronic bloaters
(122, 203); the bloater loses, by excretion,
compounds which may be essential for homeo-
stasis (203, 204).
Komarek (227) suggested that tyrosine
from legumes is decarboxylated and that its
amine derivatives, possibly tyramine, have an
effect similar to epinephrine. Leffel and
765
Komarek produced bloat in cattle and sheep
by injection of L-tyrosine in quantities that
could be consumed in green legumes (234),
but no evidence was presented to show that
such compounds are normally absorbed in ef-
fective amounts. However, Johnson and Dyer
(201) found lower mono-amine oxidase in the
livers of bloaters; this enzyme metabolizes
amines, tyrarnine being one of its normal sub-
strates, and it is generally regarded as a major
defense of the body against these compounds.
If such compounds were primary causes of
bloat, the consequence would be free-gas
bloat; they might, however, be secondary fac-
tors, increasing vulnerability to foaming.
Microbiology
Numerous attempts have been to demon-
strate differences in microbial populations in
bloating and nonbloating animals. The subject
was not reviewed by Cole and Boda (81) but
was discussed by Hungate (175).
If there are microbial differences between
animals that are bloating and those not bloat-
ing, or between animals of different suscepti-
bility to bloat, then the differences are not re-
flected in the end-products of metabolism.
Gases (176, 223), volatile acids (75, 176),
and nitrogenous compounds (75, 260) are not
different in such animals. There is also no dif-
ference in the pH of the rumen contents (260,
328).
Differences in fermentation rates between
animals of differing susceptibility have not
been demonstrated conclusively. Although
Hungate et al. (176) found higher fermenta-
tion rates in bloating animals than in non-
bloating animals, they concluded the differ-
ence was not correlated with bloat. A recent
in vitro study of the fermentation of red
clover by animals of different bloat suscepti-
bility also failed to demonstrate differences
that could be correlated with bloat (75). Ex-
cess gas production also is not proven as a
cause of bloat (175, 176), and there is little
evidence for excessive acid production being
important as suggested by Shinozaki (349).
Hungate (175) points out that legumes con-
tain so much protein and polyttronides, with
their alkalinizing effect during fermentation,
that microbial conversions would not tend to
form excess acids.
Bryant and colleagues (50) studied bacteria
in clover-fed animals and concluded that the
occurrence of bloat was not related to large
differences in the numbers or species of
ruminal microorganisms. Similar conclusions
have been reached by other workers (70, 75,
JOURNAL OF DAIRY SCIENfi]~ VOL. 57, N o . 7
766 CLAmm AND REID
157). The only major bacterial difference so
far demonstrated in bloaters and nonbloaters
is the higher number of mucinolytic bacteria
demonstrated in LS animals by Bartley and
colleagues (281, 282). Recent work in our
laboratory also supports this finding.
Oxford (298) suggested that the rurnen
ciliate Epidinium might be important in
bloat because it ingested chloroplasts contain-
ing antifoaming lipid, but an investigation of
protozoal populations in bloating and non-
bloating cattle failed to demonstrate differ-
ences (70). Mah and Hungate (245) showed
that Ophryoscolex, a ciliate related to Epidini-
urn, was in greater concentration in bloating
rather than nonbloating cattle, but the greatest
number was only 1740/ml.
Holotrich ciliates may burst following exces-
sive starch storage if substrate is in excess.
That this happens during metabolism of the
abundant starch and soluble sugars in suc-
culent legumes was demonstrated by Clarke
(71), who proposed that the cell contents
liberated by bursting would contribute sub-
stantially to foaming in the rumen. Jones
(211) has since shown that the particle-free
cell contents of rumen ciliates do produce rigid
foams over the range pH 5.5 to 6.5. Recent
work by Clarke (unpublished results) has
shown that the starch storage grains which are
released into the rumen fluid when the ciliates
burst are potent stabilizers of rumen foams.
Several trials with an antiprotozoal agent,
1,2-dimethyl-5-nitroimidazole (Dimetridazole),
were attempts to reduce bloat by elimi-
nating holotrich eiliates from the rumen
(72, 73, 74, 77, 79). Defaunation was
achieved, but bloat still occurred. The conclu-
sions were that although holotrichs are not
necessary for bloat, both the incidence and
severity of bloat may be increased in theft
presence. This finding seems to negate any
possibility of using an antiprotozoal agent for
bloat control. In a recent paper, dioctyl sodium
suffosuccinate, another antiprotozoal agent,
was effective in reducing bloat in clover-fed
steers (103) but apparently acted by slowing
fermentation. A large range of compounds was
tested for activity agains t rumen holotriehs by
Willard and Kodras (400).
Excess production of polysaceharide slime
was proposed as a possible cause of bloat by
Hungate et al. (176) and (in feedlot bloat al-
so) by ]acobson and Lindahl (184). A study
of the dissimilation of alfalfa saponins showed
that bacteria eaoable of degrading saponins,
mainly Butyrivibrio fibrisolvens and Bac-
teroides ruminicola, often produce copious
JOURNAL OF DAIRY SCIENCI~ VOL. 57, NO. 7
quantities of slime (158, 159), and it was sug-
gested that such slime production is a signifi-
cant factor in the bloat syndrome. Further
studies (157) on saponin-digesting Butyri-
vibrio showed that the slime was not the
same as that postulated as being important in
feedlot bloat (160). The feedlot slime con-
tained nucleic acid as DNA whereas this slime
contained only RNA (162). It was suggested
that in alfalfa bloat the slime is of plant, not
microbial, origin and arises from the sapogenin
portion of the saponin molecule which is left
after the carbohydrate moiety is utilized by the
bacteria (157, 162). A suggestion thai dextran
slime production by Streptococcus bovis might
be important in bloat (15) was considered un-
likely when an active dextran-splitting organ-
ism, Lactobacillus bifidus, was isolated from
the rumen in significant numbers (69).
Nevertheless, Streptococcus boris may still be
important because of its mucinolytie activity
(279).
Antibiotics may prevent bloat by slowing
gas production or by inhibiting other bacterial
processes in foam formation. Into this latter
category fall the destruction of protein before
foaming and after it becomes denatured in the
foam (75), the destruction or modification of
salivary mucoprotein and mucin (281, 282),
and of lipid (14, 247, 248, 341, 410). Penicil-
lin has affected organisms in protein degrada-
tion in the rumen (248, 356) and has been im-
plicated in lipid metabolism (248, 410). Hart-
man et al. (167) and Wiseman et al. (403)
studied the resistance of rumen facultative
anaerobes to penicillin administration. E1 Ak-
kad and Hobson (5) and Fulghum et al. (148,
391) have investigated the effects of a num-
ber of antibiotics on a range of rumen bac-
teria.
Prevention and Treatment
Prevention. Five main methods have been
used in the past 15 yr for the prevention or re-
duction of bloating: pasture management and
grazing control, feeding of roughage supple-
ments, administration of antibacterials, and ad-
ministration of so-called antifoaming agents.
Currently, prevention is based almost entirely
on pasture management and on the use of anti-
foaming agents.
Grazing legumes is always accompanied by
risk of bloat. The simplest method of bloat
prevention, complete avoidance of these
pastures, is not possible in many circumstances
and is probably undesirable if high productivi-
ty of plant and animal is to be maintained.
Control of bloat by pasture management is
 R E V I E W : FOAMY B L O A T
based on avoidance of the legume-dominant
phase in the grass sward (6, 311, 327, 345,
405). This calls for considerable skill on the
part of the farmer to control the dynamic bal-
ance between pasture species. Furthermore
limitations arise, however, from variations in
terrain in a given field and, overriding all, from
the selective grazing behavior of the animal
(318, 327).
Roughage supplements, e.g., sudan grass, ap-
pear of limited value as a practical method of
prevention (82, 83, 86, 89, 108, 169, 190,
397).
Antibacterials, the use of which followed
the discovery that penicillin could control bloat
(16, 18), are not now used extensively for
legume bloat although many papers on their
application have appeared (45, 127, 129, 130,
134, 135, 143, 145, 187, 200, 228, 256, 271,
277, 308, 309, 347, 348, 382, 383). Their fall
from favor resulted mainly from the ready de-
velopment of bacterial resistance after con-
tinuous use (175, 248, 346, 348, 361, 403).
Antiprotozoal agents have not been satisfactory
for controlling bloat (74, 77, 79, 103).
As bloat is a problem of foaming, it is logi-
cal to use antifoaming agents to prevent or
treat it. A wide variety of materials with foam-
preventing or foam-breaking activity has been
used successfully, including animal, vegetable,
and mineral otis, fats, turpentine, synthetic
detergents, and other surface-active com-
pounds (48, 61, 65, 87, 138, 187, 197, 198,
199, 262, 277, 319, 326, 330, 331, 384).
However, most silicones and some detergents
have been ineffective (330, 331).
Adequate protection from bloat can be ob-
tained only ff an effective concentration of the
bloat-preventing agent is in the rumen
throughout the danger period (319, 327). This
must be achieved in the face of the continual
turnover of contents. In lactating cows at
pasture, we have found that dilution rates for
water-soluble materials, due to water turnover
alone, may be as high as .3 h-l; ff sequestra-
tion or inactivation occurs as well, the effective
dilution rate will be correspondingly higher.
To ensure that an agent is not diluted beyond
its minimum effective concentration, continual
dosing is required. The rate of dilution of ma-
terial in solution is essentially logarithmic so
that the amount required to provide protection
between dosings increases sharply as the inter-
val between dosings increases. The practical
solution, i.e., the method of administration,
will depend on properties of the bloat-prevent-
ing agent and on husbandry of the animals.
Fats and oils lose their effectiveness quickly
767
in the rumen, and large doses are needed to
provide protection for 6 h (330). However,
by applying these materials to the pasture so
that the animal has an almost continuous in-
take of antffoaming agent, excellent control
in the field has been achieved with only
small amounts (about 90 ml off/cow per
24 h) (37, 46, 169, 180, 317, 318, 319, 326,
344). This method, pasture spraying, is still
in use in New Zealand (295) and Australia
(10) but is losing ground to methods subse-
quently described. It is well-suited to intensive
grazing where animals can be controlled with
an electric fence but is impractical in extended
grazing situations.
An alternative to pasture spraying is to add
the agent to the water in drinking troughs.
This system is less reliable than pasture-spray-
ing because drinking patterns vary with the in-
dividual and are affected by factors such as
weather, dry matter intake, and dry matter
content of the feed (58, 59, 60). A further
limitation is that animals should not have ac-
cess to alternative sources of untreated water
such as streams and ponds. Both plain and
emulsified oils have been tried in drinking
troughs (48, 283, 320, 357) but with only
moderate success, the limiting factor probably
being the length of time between drinks.
Otis also may be administered absorbed on
supplement, usually meal; such treated supple-
ments provide only short-lived protection
(191, 344), in this way being similar to
drenches (330). Another method, originating
in Australia and still widely used in New
Zealand (295), is to paint or squirt oil direct-
ly onto the flank of the animal (102, 139, 326,
344, 352), which doses itself by licking off the
agent. In both systems and also in the case oI
administration via the drinking water, relia-
bffity is basically determined by animal be-
havior. All three systems can help control mild
outbreaks of bloat.
The synthetic surfactants first tested as bloat
prophylactics were generally disappointing in
their performance (330). However, the dis-
covery that certain low-foam detergents (mem-
bers of the polyoxypropylene-polyoxyethylene
(PPE) block polymer series, "Phironics",
Wyandotte Chemical Corporation, Wyandotte,
Michigan) are highly effective for treatment
and prevention of bloat (303, 331) led to a
revolution in bloat-control measures. A PPE
polymer, poloxalene, has been developed for
use in the United States (22, 23). Other sur-
factants also have been effective, e.g., the
nonyl phenol-ethoxylate, Marlophene 89
(Chemische Werke Hills; 326, 385). In a sys-
JOURNAL OF DAIRY SCIENCE VOL. 57, NO. 7
768 CLARKE A N D R E I D
tematic search for more effective surfactants
in Australia by Laby (R. H. Laby, personal
communication) some polyethoxy alkyl ethers
(alcohol ethoxylates) were among the most
promising. These are biodegradable sur-
factants, of general formula RO(CH2CH20)-
nil, where R is a straight-chain alcohol con-
taining 12 to 17 C atoms and n is 9 to 30.
The Pluronics are water-soluble, nonionic,
and not degraded in the tureen. Not all mem-
bers of the series are useful for bloat preven-
tion. In preliminary trials in New Zealand,
some, such as the solid F68, were ineffective
as prophylactics; however, the best - the
liquids L61 and L62 -- proved superior to any
of the materials then employed for bloat con-
trol (331).
The useful polymers remain active in high
dilution; a single dose of as little as 15 g will
protect a lactating cow for 12 h or more
(192). PIurouics L61, L62, and L64 are now
established as highly effective agents for bloat
control under New Zealand and Australian
conditions (10, 192, 193, 195, 303, 304, 326,
331, 344). Regular drenching with synthetic
surfactants is the most reliable method of bloat
prevention and is gaining popularity as a rou-
tine measure on New Zealand and Australian
dairy farms (140, 141, 326, 344) when ani-
mals are handled daily in yards or stalls. A
dose of 7 to 7.5 g of a PPE polymer such as
Phironic L62, or of Marlophene 89, is given
at each milking, usually by an automatic
drenching gun (140, 141, 326); this dose is
adequate normally, but occasional failures oc-
cur when the challenge is high.
Apart from direct administration of drench-
es, Pluronics are used to treat drinking water.
The PPE polymers are better suited for use in
water troughs than are oils, not only because
of their effectiveness at low concentrations but
also because they apparently provoke more
frequent drinking (304; P. W. Young, person-
al communication). They are commonly ad-
ministered in this manner in New Zealand and,
in Australia (139, 303, 304, 326, 352), and
devices for metering them into the water sup-
ply are available. The method gives a fair
measure of control but still suffers from unreli-
ability in wet weather (326, 344; P. W.
Young, personal communication). It is valu-
able for beef cattle and dry dairy stock that
may be at pasture for long periods.
Two new methods of administration under
development in Australia (R. H. Laby, person-
al communication) are "drenching" with gelled
surfactant and the use of intraruminal sur-
factant depots. Gel drenches have the ad-
JOURNAL OF DAIRY SCIENCI~ VOL. 57, NO. 7
vantage that the protection provided by a
given dose of surfactant can be prolonged, thus
reducing the frequency of administration. The
intraruminal depots are devices which are re-
tained in the rumen for long periods and slow-
ly and continuously release surfactant at a pre-
determined rate over weeks (10, 163, 407).
In preliminary trials protection has been af-
forded for some 3 wk.
At the recommended doses, Plurouics L61,
L62, and L64 appear to have few adverse ef-
fects. Although moderately bitter, the drenches
are quickly accepted, and there are no deleteri-
ous effects on milk or cream quality or on the
properties of butterfat (195). A slight depres-
sion in milk production may occur when
Pluronic L64 is administered to cows which
are not prone to bloat (412), but when bloat
is causing a depression of milk production, ad-
ministration will prevent bloat and substantial-
ly restore production. At doses 2 to 8 times
that recommended, Pluronics L61, L62, and
L64 have caused behavioral disturbances, loss
of appetite, and scouring (in New Zealand),
and deaths have followed administration of 20
times the recommended dose (C. S. W. Reid,
unpublished results).
Poloxalene, the PPE polymer used in the
United States, is a highly successful bloat pre-
ventive and is the only antifoaming agent ap-
proved by FDA for the prevention of legume
bloat. In trials in the U.S. (31) with bloat-
susceptible identical twin cattle, it was more
effective for bloat prevention than Pluronics
L42, L53, L61, L62, L63, L64, L92, L101,
P105, and F108, particularly at low doses (2.5
g/454 kg body weight). In studies with 120
lactating cows, poloxalene was more palatable
than Pluronics IA2, 1_,53, L~61, L62, and L92.
Pluronic L101 equalled poloxalene in palata-
bility but was ineffective as an agent for bloat
prevention (31).
At the recommended doses, poloxalene has
no deleterious effects on feed intake, rumen
fermentation, animal health, weight gain, re-
production, or milk production (22, 23, 24, "38,
170, 232, 238, 241, 274, 316). Some 4% is
absorbed from the gut and appears in the urine
(269), but no residues appear in milk or tis-
sues (268, 269).
Methods of administration of poloxalene are
generally based on feed or feed supplements
as vehicles; the surfactant may be incorporated
in the ration or sprinkled over grain (27, 29,
133, 142, 144, 366, 368, 369), incorporated
into molasses salt blocks (131, 132, 142, 144,
170, 367), or offered with molasses in tanks
with lick wheels (30, 342).
 REVIEW: FOAMY BLOAT 769

may be present. Research effort is the poor re-

Synthetic surfactants affect surface tension,
but how they prevent bloat is not fully estab- lation of pasture bloat. In spite of the availabil-
lished. Poloxalene has lowered the surface ten-ity of a ration which will produce feedlot bloat
sion of rumen contents (316, 367); a single (353) consistently, progress has been slow
dose of .044 g/kg body weight reduced rumen towards discovering the cause of foaming
surface tension for 24-36 h (316). Aqueous (66). It is difficult to separate the literature
solutions (.001%) of L61, an effective on feedlot and pasture bloat. Some papers do
Pluronic, have a lower surface tension than not distinguish the diet of the animals and in-
those of F68, a less effective polymer (331). terchange freely the symptoms and effects of
Laby (R. H. Laby, perso~aal communication, each type of bloat.
406) has examined a wide range of sur- Feedlot bloat traditionally develops in cat-
tle when concentrates form 50% or more of the
factants, including poloxalene, seeking to relate
physical properties with efficacy in bloat pre-ration fed (183). Unlike pasture bloat, feedlot
vention. He has found that detergency and bloat usually develops slowly over weeks and
wetting power correlate better with success in often becomes chronic. Almost all of the tradi-
bloat prophylaxis than do foaming properties. tional diets may give rise to it, but the most
serious bloat generally occurs on the finishing
The long persistence of the effects of synthetic
surfactants might be explained by adsorption ration. Bloat arising from the starter ration may
and subsequent slow release from the absorp- be caused partly by acid indigestion.
tion sites. That a large part of a dose of sur- The role of roughage in feedlot bloat is not
factant is associated with solids in the rumen clear. Although roughage is accepted general-
contents supports this hypothesis (R. H. Laby,ly as reducing the degree of foamy bloat, alfal-
personal communication), as would the low fa hay is considered by feedlot operators to be
concentrations, but apparent slow dilution, of a bloat-provocative roughage (E. E. Bardey,
the surfactant dissolved in the liquid fractionpersonal communication). Alfalfa meal is a
(Reid, C. S. W., and D. R. Body, unpublished common constituent of feedlot rations and
comprises 22% of the experimental bloat-
results). An intriguing possibility is that these
inducing ration of Smith et al. (353). Why al-
materials may influence the distribution of fats
falfa hay differs from other roughages has yet
within the digesta; synthetic surfactants could,
in effect, recruit the dietary fats, releasing to be determined. Gordon and Erwin (151)
them from sequestration sites and allowing found bloat occurred only when alfalfa hay
them to become effective as antifoaming (30%) was added to the basic cracked corn,
agents. soybean meal diet of their steers. Phillips
Treatment. The administration of an anti- (305) (quoted by Preston, 314) observed
foaming agent remains the best method of bloat in his animals (calves) only when they
gained access to hay, wood-shavings, or leaves.
treating cows suffering from pasture bloat. Ani-
mal, vegetable, and mineral oils are effective On the other hand, grinding the diet has in-
(189, 192, 199, 326, 330). The polyoxy- creased the incidence and severity of bloat
propylene-polyoxyethylene polymers including (84, 91, 171, 259, 314). The possible modes
poloxalene (29, 32, 326, 331), phenol of action were discussed earlier in this review.
ethoxylates (326), and the alcohol ethoxylates However, some features reported by Hironaka
are highly effective; they are the materials ofet al. (171), such as the rapid onset and the
choice because they relieve bloated animals low pH in rumen contents, suggest that mild
more rapidly than do oils and have the further degrees of indigestion also may be involved.
advantage of providing protection for the fol- The occurrence of feedlot bloat is not re-
lowing 12 h or more (331). Direct injection lated to eating habits, nor is its severity related
of antifoaming agent into the rumen is not as to increased food consumption (236). Volatile
satisfactory as drenching (29). The use of a fatty acids are not different in bloaters and
stomach tube may be of value for individual nonbloaters (124, 126, 186). The pH of the
cows to release any free gas but is not usuallyrumen contents of bloated animals is not gen-
necessary. In acute cases, particularly when erally low (125, 126, 161) and does not ap-
animals have collapsed, rumenotomy may be pear to be a significant factor (1"25) except
essential to save life (169, 192, 320). where acid indigestion occurs as well (51).
Emery et al. (128) examined various com-
FEEDLOT BLOAT ponents of saliva and tureen fluid of two cows,
Feedlot bloat, like pasture bloat, results a bloater and nonbloater. The differences in
from excessive foaming of the contents of the saliva were only in Na, K, and P and did not
rumen (185, 236, 263). Some free gas also appear large enough for them to be important.
JOURNAL OF DAIRY SCIENCE VOL. 57, N O . 7
770 CLARKE AND REID
Increased amounts of dry matter, organic mat-
ter, Na, hexosamine, total N, and 81~o
ethanol-soluble N were in rumen fluid of the
bloater. However, such differences could have
represented individual animal variation, per-
haps related to intake but not necessarily to
susceptibility to feedlot bloat. Higher
hexosamine and nitrogenous compounds could
have been partly a reflection of salivary mucin
and mucoprotein. Jacobson and colleagues
could not demonstrate a correlation between
dry matter content or consistency of rumen
contents and bloat (184, 185).
The foam in feedlot bloat is thought to be
microbial in origin. An ethanol-precipitable
slime is in the rumen digesta (160), and its
concentration correlates loosely with oc-
currence of bloat. The slime was composed of
34% crude protein, acid-hydrolysable poly-
saceharidcs, and nucleic acids (160). In con-
trast with the slime from clover-fed cattle
which contains RNA, the slime from feedlot
bloat animals contained mainly DNA, indicat-
ing that it was probably of microbial rather
than plant origin. Differences in microl~ial pop-
ulations of bloated and nonbloated animals
have been reported. The numbers of encap-
sulated bacteria increase in bloating animals
(185), and increases in encapsulated Strep-
tococcus bovis and Peptostreptococcus elsdenii
have been reported (161, 278, 279). However,
S. bovis may become reduced in numbers after
long adaptation to the diet (51).
Bacterial capsular material may increase
viscosity of rumen fluid. Most streptococci ap-
parently associated with feedlot bloat are
thought to be Lancei~eld Group D (263), the
members of which have rhamnose present in
their capsules (172). Meyer and Bartley (263)
made a preliminary attempt to relate feedlot
bloat to viscosity of rumen fluid and to extra-
cellular polysaccharide (glucose and rham-
nose) but concluded that further work was
necessary before a correlation could be estab-
lished. Bryant et al. (51) found great variation
in microbial populations of two steers bloating
on the same ration. They concluded that feed-
lot bloat can occur in animals with widely
differing numbers of any of the individual
groups of bacteria. Mishra (280) found proto-
zoal popul, ations were reduced in bloating ani-
mals.
Kodras (226) suggested that gas bubbles
expelled by entodiniomorph ciliates may be the
cause of bloat. It is our experience that ciliates
expel gas bubbles only when surrounding
medium is conducive to bubble formation. The
phenomenon seen by Kodras probably indi-
IOURNAL OF DAIRY SCIENCE VOL. 57, NO. 7
cated that the bloating rumen fluid had differ-
ent surface properties to, or was more viscous
(e.g., had a higher content of slime) than,
nonbloating fluid.
Salivary mucin and mucroprotein possibly
contribute to foam formation in feedlot bloat
as they have done in pasture bloat. However,
there is no direct evidence to show this. The
foam-preventing activities demonstrated by
Bartley and colleagues also may operate: the
addition of several mucinolytic bacteria to the
rumen caused the onset of feedlot bloat (136,
279).
Feedlot bloat in beef and dairy cattle and
sheep can now be prevented by oral adminis-
tration of a dimethyl dialkyl quaternary am-
monium compound (265).
Although the polyoxyethylene poly-
oxypropylene block polymers are effective in
dispersing foam in pasture bloat, they are not
as effective for the treatment and prevention
of feedlot bloat. However, poloxalene can be
used to prevent bloat in cattle feeding on alfal-
fa hay and barley (149). Use of poloxalene on
severely bloated animals does not cure the
complaint but does free some gas which may
be removed with a stomach tube (E. E. Bart-
Icy, personal communication, 28). Mineral oil
is satisfactory in concentrations of 4 to 8% of
the ration, reducing bloat by 40% (124), but
animal fats are not effective. Soybean oil fed
to cattle as part (8%) of the ration resulted in
a highly significant increase in bloat (124,
126). This difference in behavior of mineral
and plant oils may reflect a role for lipids in
the formation of feedlot foams. Other com-
pounds, including various surfactants, sili-
cones, fatty acids, antibiotics, and dyes, have
not proved successful for control (264).
Although pasture bloat may be a conse-
quence of ingestion of a diet more succulent
and richer in protein than the ruminant has
evolved to cope with, the feeding conditions
in feedlot bloat are in this sense, even more
abnormal. The low fiber content of diets induc-
ing feedlot bloat will have effects on salivation,
rumen movements, and rumination, all of
which are likely to be reduced markedly. What
contribution such effects make to susceptibility
is not known.
CONCLUSIONS
Bloat is still a common and costly disor-
der. An integrated picture is starting to emerge
after more than 30 yr research, but much re-
mains to be done, especially for feedlot bloat,
before the complex interactions are unraveled,
Such detailed studies will help the develop-
 R E V I E W : FOAMY B L O A T 771

ment of long-term prevention measures and rumen of salivary components, especially

possibly lead to elimination of the problem. As mucoproteins and mucins, the amounts and
well, bloat research will continue to provide composition of saliva secreted during bloat-
fundamental knowledge of wider application. ing, the 24-h secretion pattern and its influence
Below are set out some of the areas in which on susceptibility. The possibility should be
we believe research should be concentrated. considered that properties of saliva other than
Before considering them we stress again that those already discussed, such as the presence
more care should b e taken to define the cir- of antibodies which might affect the microbial
cumstances and degree of bloating, as well as population, also may be important.
the innate susceptibility of the experimental Particulate matter. Particulate matter, rang-
animals (78). Palpation of the left flank (189, ing from bacteria to "coarse" plant fragments,
199) remains the most reliable of the simple is a characteristic feature of namen foams. Lit-
methods of determining the severity of bloat; tie is known about the dynamics of the particu-
it can be related accurately to intrarnminal late fractions in lumen contents - the mecha-
pressure by an experienced assessor (240). nisms and rates of eomminution of the feed,
The foam. The actual components of the the range of particles that results, the distribu-
foams, their roles, and associations have yet to tion patterns of .the finer particles, including
be settled. Until these are known, it will not the effects of foam flotation, the rates of exit
be possible to detect variations in proportions of the different sized fragments, and the fac-
of the components, or possible substitutions, tors which control their outflow. Another
and correlate these with variations in potency aspect of particulate matter is the tactile stimu-
of feed or severity of the disorder. lation afforded to the sensory nerve endings in
On the physical side, data such as the tem- the stomach walls and its effects on salivation,
perature of the rurnen contents at the time of rumination and rnmen movements, and eructa-
foaming are still required (52). Information tion.
is also needed about surface properties of the Gas elimination. Bloat results from a shift
natural surface-active agents in the rumen, in the balance of gas production and gas elim-
both individually and in combination. How- ination, yet little quantitative data are avail-
ever, more realistic simulations also are able on gas dynamics even in the normal, un-
needed. Simple in vitro foaming systems ignore bloated animal. Specific areas warranting fur-
normal chemical, physical, and biological com- ther study are the manner of interference with
ponents. Work is now required in which elec- the eructation of free gas, the reflex effects of
trolytes, surface-active agents, particulate mat- the entry of foamy digesta into the esophagus,
ter, active microorganisms, and the like are and the possible interactions between foaming
added and the experiments carried out under and impaired gastric motility.
the appropriate conditions of temperature, Consequences of gastric distention. Al-
pressure, osmolality, and gas composition. The though a number of studies have now been
dangers of extrapolation from in vitro to in on physiological effects of distention of the
vivo situations have been emphasized by rumen, the circumstances of the studies have
Dougherty (78). In the animal itself, attempts been more akin to free-gas bloat than to pas-
could be made to modify conditions within the ture bloat. Further information is needed about
rulnen. the consequences of ruminal distention in
Lipids. A gap in knowledge is that of the bloating animals and its effects on circulation,
role of lipids in bloat foams. Their presence in salivation, gastrie motility, CO~ absorption, and
large amounts in both legume and feedlot acid secretion in the abomasum. The problem
bloat foams has been confirmed by several of cardiospasm also requires examination to es-
workers, but whether they are acting as anti- tablish the exact nature of the phenomenon.
foaming agents, assisting protein binding, or Surfactants as bloat-preventing agents. Al-
are playing some other role, is not known. De- though there is no doubt as to the effectiveness
tailed information could provide explanations of the synthetic surfactants to prevent pasture
for the short-lived protection by oil drenches bloat, little is known of the details of their op-
and why a nominal 80 g of vegetable or eration. The distribution of the surfactants in
paraffin oil will as a pasture spray protect an the rumen, their effective dilution rates, and
animal for 24 h when the normal daily intake their interactions with lipids and other natural
of pasture lipids is Eve or more times tl~at surfactants all need further study.
amount. Tannins and the breeding of less-bloaty
Saliva. Aspects of saliva on which more in- legumes. Research has been focused on the
formation is needed include the fate in the role of condensed tannins in precipitating plant
JOURNAL OF DAIRy SCIENCE VOL. 57, NO. 7
772 CLARKE AND REID
proteins during chewing; however, as they will
precipitate salivary proteins in vitro, it is pos-
sible they also have an effect on these pro-
teins. The fate and effects of tannins in the di-
gestive tract are not known.
T h e breeding of clovers containing increased
condensed tannins is under study at this
laboratory. Several years work may b e re-
quired merely to establish the feasibility of the
scheme, for as well as containing increased
amounts of tannin, the new plant must also be
agronomically and nutritionally acceptable.
Nevertheless, it appears a more acceptable
solution biologically than breeding plants with
lower leaf proteins.
Susceptibility and breeding animals with re-
duced susceptibility to bloat. T h e factors con-
trolling susceptibility, the manifestations of
the trait, and its stability all need definition.
The inheritance of susceptibility must b e de-
termined, both in terms of degree of suscepti-
bility and of the controlling factors.
The breeding of cattle having reduced sus-
ceptibility to bloat is possible in theory.
Whether useful changes can be achieved in
practice will depend on factors such as stabili-
ty of the trait, complexity of its inheritance,
and rate of decrease in susceptibility through
successive generations. Progress is likely to b e
slow unless a marker that correlates with sus-
ceptibility can be found.
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