Ondetsfepoort .!outnal of 1-r,fctinary Sr·ie11ce and Animal lwlusfT'!}, l-ol11'1ne 2:1, .\'·11mhen 1mHZ2, Jllanh, 1948.
1Pri11ted in the lJ11io11 of South Afri('a by tile Go,·erllll1E'IIt l'ri11ter, Pretoria.
Studies on the Alimentary Tract of Merino Sheep in South Africa. XIV.-The effect of some commonly used Antifermentatives on the in vitro Formation of Gas in Ruminal Ingesta and its bearing on the Pathogenesis of Bloat.
Jh H. ('LAHK, St•dion of J>h~·siolog_v, ()IHierstepoort..
I~TitODUC'J'lOJ\.
0 1\.L\'G to more intPnsiYc fanning 111ethoch <llld the inneasiug use of lucerne a~ a fee<l, till' problem of hlo;tt in ruminants is assuming g-reat importance in f-:lout lt A hie a.
Although this cmHlition has heen kno\l'lt th<• "·orlcl o1·cr for cenituie~
its adunl pathogene,.;1,; i,; Jtot yet t·learl,v untlf'rstootl. J n a rt'CPnt exhau,.;tin•
review of the t>uh.ied by Cole, Hnffm<ln, Klei1wr, Olson antl Schalk (194!)) it is sbtt:><l :-
" A YiP\\' h<·ld until n•centh I\' a,; thai. onl 1· uu<h•r conditions in ,,·hiclt l1loat \\·as t·outmon wa,.; thl'l'f' a11 appreci'able formation of ga:-;
in the rumen. l<'nnn eYi(h·n<·e Jt011· at han<l, llOI\'PYE'l·, tlw fee<ling of bloat proYoking tlieb tloes not nC'l'e,.;saril~- n·sult. in gn•atPr g-as forma- tion- '' aiHl later "- t lw main proble111 in bloat S<'Pllls to !'t>ntl·e arOUllll till' e:xpubion of g·a,; !'athn tltnn ih formation."
Blo;d. is oln·iousl~· the n•sult. of g·as 1Jl'ing- fonlletl quil'ker than it is elintiuatr•d. ('lark and quin (194!)) ll<t\'e sh<mn ihat sheep ('au eructate ga,;
(CO" or Ail') <tl'tifi('iall:· inhotlucetl into the nUJtPn at t hf' rate of bYo litrb per minute. As this i;:; undo1tbte<ll.Y far in excess of the possiblt:> gas evolu- tion in the rumen, thE' <tl'I'Umnlation in bloat mmd be as:-;oci<~i.ecl with bilun·
to Pliminate i1 sufficiently rapiclly.
The time-honoured awl undoubterlly sU<Tessful use of turpentine as an antifermentatiYt- in tltt:> ht><~huent of t.:nupanites rloes lint appear
to
cm1form wit It tlw n hoYt> YLe\\·. H 11·as ih erefoJ'e <lel'ide(l to ascerbt in at IYha t con(·pntra- tions some of the more t·ominonly athoc:atP<l rlrug,.; in II ihitt>tl ga,; formation in rumi11al ingesta.}!ET!LOD.
8i.x merino ,.;heep ,,-ith permanent nuninal fistulae were usecl, lucerne- hay ud lib. hE'ing- fe,l PxdusiYeh·. Samples of nunen ingesta were strained through butter muslin alHl 20 , .. c. nliqnob of thP flui,l placecl in test tnhes.
Glucose was aclded to nll tuhf's to bring about a final eon('entratiou of 0·8 per Cf'nt. Eight ~u('h tubes hom en,·h of the six sliE'ep were sE't up at a time. 'rhe first hYo of each set aderl as controb and the agenb; to be teste,l ll'ere aclded in tl1P rlesired ('oncentrations to the remai.nJer, duplicates being used fm ea('h treatment. After Yigorous shaking Dtnham fermenbtio11 tubes ''"ere inverted in the te;;t tubf's al!ll the samples incubated in a water bath at :i9° C. for tw-o hour;;. The length of the gas c·olumns was then measured in millimeterfi.
The ave1·agp results of nmnerous sn('h triab are g-1Yen m 'ruble 1.
Drug.
'.L'urpentine ... .
Copper Sulphate ........... .
T.\HLE 1.
Therapeutic Dose for f>heep.
*
l :i C'r.
1 gm.
C:t1cu1a.tecl Concentration
in Rumen.
t
1,000
I : I :),000
Concentration Tested.
1,000 +00 200 100 1.),000 10,000 : -!,000
Percentage Reduction in Gas.
() :I 3:1
8;)
!l (i~
!)2
--- i---1---
: 4,000 I :i
: 2.000 :~()
: 1;ooo 70
Foema1in ........... . -i CC'. I : :1,750
- -- -- - - - --- - - - ; - - - 1 -
1:),000 : 1,000 4
: 400 12
Phenol. ...•.............. 1 gm.
: 200 -i!)
- - - -
: 4,000 0
2,000 17
1,000 uo
Peoprietary Coal Tar: Prep. A.t
I
-i cc. 3,750______ ________ ___ ! _______________ - - - · -
4.000 (i
Peopeictary Coal Tar: i>rep. H.t
I
4 cc.I
l : 3,7:i0I
-~---
*The maximum doses found in standard text books.
t Calcula.ted on a sheep rumen of I :),000 cc.
t These >tre both peoprietary dips or d.isinfcct:tnts containing phenoloids (HP. 200-300°C) and coal tar hydrocarbons largely nsed by farmCI'R in 1-'outh Africa. for the treatment of bloat. The recom- mended dose of both for sheep i one dra.chm.
Considerable Yariations in the perl'entage ,le,'l'e<lSe in g"<1s procluction witl1 giYen roll(·enhationfi of t·he ,litY!:'rent agent;; W('l'e fouu(l nmong- thP sample<; hom the nni.ous animals. The thic·ker and more vi,;cid the ing-e;;ta the le:ss was g'<Js procludion afTPde(l by tlw di;;infedant. Thi;; '"a,; rlouhtles:;
390
f.
R. CLARK.
due to the inhibitary action of the orgauie matter on the agent. The follow- ing extracts hom the uata illustrate this point.
Date of Collection
I
Date of Collection
28/11/45. 4/12/45.
Type
Sheep. of Gas Produced. Gas Produced.
T ngesta.
I
Control. I .Plus l : 400 Control.
I
Plus 1 : 2,000Turpentine. Prop. Prep. A.
L .•••.•••.•••••.••• Viscid. 70 68 63 54
2 ........ Viscid. 69 66 37 20
3 ... Watery. 12 0 22 8
4 .............. Viscid. 67 70 43 40
5 ... Viscid. 6ii 64 42 31
6 .................. Viscid. 50 46 53
I
48--
As the viscous, high gas producing type of ingesta is invariably found iu cases of acute bloat on g-reen feed, it is to he expected that, under these conditions, the agents would be eYen less effective than inclic:ated in Table 1.
CoxcL US IONs.
In view of the above findings it can he nssumed that neither turpentine nor the coal tar clerivntives have any effect on gas formation in the rumen in therapeutie concentration;;. Sonw other explanation must, therefore, be found for their undoubted beneficial effect in the treatment of aeute tympanites and this explanation may help to eluci(late the pathogenesis of tlte condition.
DISCCSSIOK.
'fhick Yiscid ingesta are often foamy \Ylten withdrawn. The foam breaks only slowly on ;;tatHling and refonns with shaking. The addition of small amounts of turpentine, ('oal tar preparationR or thin oil brings about a11 immediate and marked change in the physical propeTties of the ingesta, breaking the foam and preventing its refonnation. 'l'hi;, clearly indicates that it is through their aetion on surface tension that these drugs exert their beneficial effect in cases of acute hloa t. This explanation is based on the hypothe;;is that hothing of the ingesta is au important factor in the pathogenesis of the eouditiou.
This theor~r is hy no meau;; new and has been suggested by McCandlish ( 1937), Olson ( 1942) a n<l Quin ( Hl40), a 11 of wl10m incriminatrd saponin aR being the causal factor. .1 a<·.ohson (1919) fed 19 gm. of lucerne saponin to a sheep without ill-effects HJHl tbe present author has failed to cause retention of gas artificially introrlnced into the rumen h~· previous closing with saponin.
The frequent oc<·unellce of frothy ing·estn in as,;ociatiou with acute bloat, accompanied by failme to relieve the eoll(lition hy pnssage of the stomach tube or insertion of a, canula, is, however, well lotown to all who have had experierwe of the vondition. The sheep \\·ith permanent fir;tu]a,e at Onderstepoort frequently exhibit marked clistention but removal of the cork from the fistula
(!
inch diameter) often fails to relieve the pressure, only a small amount of froth f'XlHling throngh the tube.ln tht> reyjp,,- JHHiousl~- l'ited (Cole et ol, Hl45) Hufflll<lll di:-;(;u:-;,;es tl1e hotb theon· of bloat hut cloes not appear to l1e at all conYinced of its Yalidity. Ife state s : -
" Cole autl his c·o-worker,; ha1·e bppl'd a nu111lwr of c·o1Y:-; hloatt•cl on alfalfa pasture without encoun!Pring- a t·ase in IYhieh the hc>e air spal't' was abseut. The fad tlwl often no free g-us hut·, rathPr, ho!ll\.
ing-esta, esc;qws 11·hen tlH· I'Uinell is tuppc-•d is regardt'd by Jllany as proof tl1at free g-us is 110!· pn·senl. The c·ausP, c·onl'eiYahh, ma_,- lw the fad. that tl1r .irep air ,;pan• is tliffil'ult to tap 11·hen the l'111llen i,;
excessiHly filled. If a free air spa('p PXish, it is cliftic·nli- to nndPr- :-;tand ''"'''" frothing- ll"lllllcl c·unst· hloat. At the mon1eni at 11·hic·lt helchi11g· oc·c·un there is <l dilal ion of I he retieulun1 <lnd anterior l'lllllPn, thus tt>nding to hPl' !he t·ardi;t fm· the passage of ga,; into the oesopbag'JI,;,
The prt'seni <111thor ('f\nnot ag-nl(-• with tltest• c·o11clusions.
G-as i,; n•moYecl hom the rumen hy t11·o methods (i) clired absorption into the bloocl ,;hP<llll thnlllg-h tht• J'11111t'll 11·all ancl s11h-;eqnent dimination hY tbe lun).!'S and (ii) lwlc-hing.
The proportion of gas Plimin;dc·d IJ_,- the fi1·st nlPthocl is not kno11·n hut it has frequently heen clelllonstrated t·hat Llrge amon11ts of carbon dioxidt•
and nlt>lh<llle ;11·e <lhsorhecl through tl1P l'11111en and th;d the ndt• of such absorhtiun is clependent on tht• jll'l'.~~Ul'l'. The l1ean· clPPIJ breathing· Kl'Pil in llOll-hloatecl aninwls shodh- ;dtt•r fec•cling- on lnc·erne is eausecl h,· tht>
ahsorptio11 of ('arbon-clioxide fnnn the rumen, tht• S<lllll' el'fec·t being- produc·Pcl by al'(ifi(;ial illfla!ion of the nllnt·n 11·ith c·<Jrbon clioxiclt• (('l:nl; <11Hl (luin l!J45). If the g-al'i is irappecl ill " tc•1wc·iou~ foam it ''"ill11oi c·on1P into !'Oniad witl1 the ntminal lllll!'OS<l <Jllll ab~orpt-ion ''"ill be 1·eL1nlPcl.
1Tllfortull;deh tht• mec·h<tni,;m of !he erndaiion 1·eflex in J'lllllin<lllts is not dearl:· nn<le1:stood. Cole, "Jle<td and Hc•g-an (194:)) haw f<mnd <l roug·h l'onelation lwf11·een J·umiiwticlll ancl belc-hing- <IJHl ,;ug-g-est· th;d il1P st i11n!lu,;
for the hu> ads is the same, name]_,. thl' pre,;enc·c· of roug-h scabrous· mate1·ial in the rumen. This has been JHOn·c1 to he the cast• in rumilla!iott (:-:.dJ;dk and Amadon, Hl:28). Cole et of. (1D4:~) suhstantiate their hypothesi,; by !he well-kno\1"11 fad tl1at feerling- l·oug-h h<1~· before luc·erne iellcls to preYl'nt bloat hut thi;-; !'<Ill <tlso bt:• PxpLlllH'd on the foam thPor.\'. The aclcliticm of l'i10ppecl hay to s<!mple,; of 1·enclih fo::n11ing· ing-Psb breaks the froth. It is difficult to unclen;tancl wll\· rumillation <llld belching, <tlt houg-h simiL11·
met·hunie<lll~·, should both he initiatecl by the p1·es!'ll!'E' of c·ourt>e materi;!l in the n1lneu. Such matter requir!'s rPche11·ing- <lllll so it is log-il'al that ih presence Hhoulcl c·<Ju,;e rnmillatioll hut as it proditce,.; littlc• g·as its \'Olllledioll 11·ith belchillg- is obcure.
The ;t,;~oc·iation IH·tween run1inal moiilih· aucl erudatioll i,; also not
c·lear. Clark and Quill (Hl--!5) lmn• ,;ho11·n th:it unclPr c·l·rtnin cin·umsta11c·ps sheep ('all helc·h ga,.; artificiallY illtrocluc·pd illto the l'lllllt'll despite ruminal pare~is inclU('t>d hy pl'u,;sic ac·icl. C'oui·Prsl'h· hcquc·nt <11Hl ,;trong· ruminal
!'on tractions are often ,;een i11 <Jni n1<1ls suffel'illg- ho111 Ul'Utt' hlo;d.
Free g-n,.; 11·ill accumulate in the dorsal S<J!' of !he rUlll!'ll ancl in ordc·r io reach the l'<~rdia it must he lii!>Ye(1 fon1·ards uncl rlownwanls. 1'he pa,.;e
~~·ith "·hic·h this can be rlone will clt"pencl 011 tlw Yolume ancl YiS!'osit_\· of the Jng-e;;ta. \Vhile filling the Durlunn tubes in the I)l'esent experiment it· "·"s found that, in the l·ase of the thick tenaeinus samplt>s, ,·ig-onm;-; ,.;haki11g- in
392
~s the
b f
itsoated e au·
·othy
Y as
~· l~e en 1,;
nder- ,·hich fen or
into
ption ation
but xi de such seen the uced Quin utact b is nwh tuh1s erial halk
; the Yeut n of
[t i~
1ilar
"rial t it;;
·tion not nces inal inal l'cler ease
the was
~ lll
A
'"
R. CLARK.
the inverted position had to be l'esorted to in order to get the air out of the tubes. The air escaped easily in both the watery sample:S aml the thick samples after the addition of turpentine. The dosing- of turpentine will therefore not only break up the foam an<l inrrease the volume of free gas available for eructation hut also facilitatt> its b·ansportation "·ithin the nuniua l mas;;.
Cole et al. (1945) state that belching is facilitated b~r the administration of turpentine, but the exaet mode of action has 11ot :vet been determined.
They postulate that it may, by its initant action, initiate the reflex. This is a possibility that cannot he denied but it does n·ot invali<late the beneficial pffeds on surface tension already discussed.
In the w1·iter's opinion turpentine is still the best drug known for the treatment of ~H;ute bloat but as it acts largely through its effect on surface tension superior preparations ma~· yet he found. It is very doubtful whether disinfectants should he introducer! into the nnnen even if thPy are effective in 1·educing g:1s formation. A properly balanced arHl active microflora is essential to ruminal digestion and its impairment or destruction may have RPrious sequence. It is sugg-ested that in milder eases turpentine be dosed, p1·eferably through the stomach tube. In critical cases the canula should be im;ertrd and the turpentine injede<l through it. This will eause disruption t'f the foam at the end of the canula and a consequent escape of gas.
Sc);aLI.RY.
1. It has been shown tl1at turpentine and hro proprietar~r coal-tar preparations have little or no inhibitary action on g-as formation in ruminal ingesta incubated -in rih·o when added in therapeutic roncentrations.
2. Very small concentration~ of thei>e substances have a marked efted on the physical consistency of the ingesta, raising surface tension and breaking down foam.
3. This affords evidence that it is by their physical action on surface tension that these drugs exPrt their ur~iloubted bene£cial effect in acute bloat.
4. 'rhe "foam theory" of the pathogPnes.is of bloat i;; <liscutiseil.
5. :Formalin and eopper sulphate may have some inl1ihitory action on gas formation but the use of n.m1inal disinfectants cannot be recommendeil.
LITERATURE.
CLARK, R. AND QUIN, J. L. ( L94Fi). Studies on the >1limentary tract of merino sheep in South Afeica XIII. The role of prussic acid in the aetiology of acute bloat. Onderstepoort Jnl.,
Vol. 20, No. 2, pp. 209-212.
COLE, H. H., MEAD, S. W. AND REGAN, W. M. (1943). Production and prevention of bloat in cattle on alfalf<t pastur·e. Jnl. Animal Sc., Vol .2, pp. 21\ii-294.
COLE, H. H., HUFFMAN, C. :F., KLI~IBER, M. OLSON, 1'. :M. AND SCHALK, A. F. (194ii).
A review of bloat in ruminants. Jnl. Animal Sci., Vol. 4, No. 3, pp. 183-236.
JACOBSON, C. A. (1919). Alfalfa saponin. Alfalfa investigations VII. Am. Chern. Soc.
Jnl., Vol. 41, Part l, pp. 640-648.
McCANDLISH, A. C. (1937). Hoven or bloat in dairy cattle X f. Milch11·irtschajtlicher Weltkongress, Berlin, 1 : pp. 410-412.
OLSEN, T. M. ( 1942). Rumina! gases in normal and bloated animals. Jnl. Dairy Sci., Vol. 25, pp. 684-685.
QUIN, J. I. (1943). Studies on the alimentary tract of met·ino sheep in South Africa VIII.
The pathogenesis of <tcute tympanites (bloat). Onderstepoort Jnl., Vol. 18, pp. 113-118.
SCHALK, A. F. AND AMADON, R. S. (1928). Physiology of the ruminant stomach (Bovine).
Study of the dynamic factors. N. Dak. Agric. Exp. Sta. Bull., No. 216, pp. 1-64.
