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Piis0022030291785512 PDF
Piis0022030291785512 PDF
size also is involved in the rumen buffering P-glucanase, and protease (25). In some of the
system along with nonstructural carbohydrates modified methods, the sample is incubated
(NSC)and fiber quality. overnight at 20 to 35'C with amylase. With
this longer incubation time, contaminating en-
METHODS FOR NDF zymes in the impure amylase preparations can
degrade hemicellulosic components in the
The insoluble fiber in feed includes the feed, giving low values of NDF, whereas in-
crosslinked matrix of the plant cell wall and, as clusion of unwanted starch leads to high values
coarse fiber, forms the rumen mat that stimu- (25). Enzymatic preparations from Bacillus
lates rumen function. It is measured most con- subtilis have been most popular. However,
veniently as NDF, which includes cellulose, their quality and availability has varied over
hemicellulose, and lignin as the major compo- the recent yeaxs. The amylase (Sigma Type XI
nents. The original NDF method was applied is no longer available) contained sufficient
to forages, and its subsequent application to hemicellulase to affect values of wheat bran
starchy foods and feeds revealed interfemce and possibly wheat straw (Table 3). Avoidance
by starch, thus presenting difficulties for the of this problem was accomplished by conduct-
original neutral detergent (ND)method. There- ing starch digestion at the highest possible
fore, various modifcations with amylases have temperature. A newer B. subtilis enzyme
been reported (Tables 1 and 2). Many of the presently available from Sigma is of lower
commercial amylases that have been used con- activity, and more enzyme has to be used. The
tain other activities, including hemicellulase, B. subtilis amylase is limited to a temperature
Journal of Dairy Science Vol. 74. No. 10, 1991
SYMPOSIUM: CARBOHYDRATE MeTHODOLOGY 3585
TABLE 2. Description of two neutral-detergent (ND) methods using differing Operating conditions and two amylases.
Reference Conditions AlXOUpl
Jeraci et al. (17) Single addition of heat stable &amylase to ND solution, boil 60 ND-T
min
Robertson and Van Soest (36) Boil sample for 50 ml of ND solution for 30 min and then add ND-S
50 ml of ND solution and Bucillus subtifus a-amylase, M i 30
miq filter and add B. sltdrilur &amylase to crucible, incubate
for 10 to 15 min
of 80°C and is rapidly inactivated at 1OO'C. were effective in removing starch (16). Al-
The a l - 6 activity also is destroyed by EDTA though wheat bran gave less NDF by the ND
because it is calcium-sensitive. In some cases, assay using the B. subtilis amylase, the starch
the enzyme was added during and after the content does not account for the difference
refluxing step to remove the starch (36). These (Table 3). The differences between NDF
modified methods increased the assay time by values for the samples cannot be attributed to
only 5 to 20 min but increased the number of starch; they probably reflect a loss in hemicel-
steps in the ND procedure. lulose, as also noted by Mascarenhas-Ferreira
This new heat stable a-amylase is stable to et al. (25), who found even larger losses in
boiling detergent and is used to degrade starch hemicellulose with lower temperature treat-
in nondetergent chemical methods for dietary ments.
fiber (19, 35). This a-amylase, which has In the original ND method, starch removal
AOAC approval (Number A3306 in the dietary was facilitated by using 2ethoxyethanol. How-
fiber kit; Sigma Chemical Co., St. Louis, MO), ever, Z-ethoxyethanol (ethyleneglycol mono-
has been used effectively in the ND method ethyl ether or cellosolve) now is recognized as
(16). The use of high temperature with a short- a health risk Its use appears necessary for
term amylase treatment has the advantages of optimal removal of starch (16). Therefore,
minimizing the effect of unwanted side activ- 2ethoxyethanol should be replaced by a safer
ities and is a more rapid procedure. reagent. Use of triethylene glycol at the same
We have compared ND procedures using concentration gives equivalent values and is on
either B. subtilis amylase (36) or heat stable the safe list. Thus, even with the use of effi-
amylases (Tables 2 and 3). Both amylases cient amylases, addition of either 2ethoxy-
TABLE 3. Evaluation of two amylases in Iwo neutral detergent (ND) methods. Analysis of NDF and the content of starch
in NDF.'
ND-S2 ND-9
Sample NDF SEM4 Starch NDF SEM Starch
(96 DM)
Timothy hay 65.5 1.31 0 66.3 98 0
Wheat straw 83.8 .ll 0 85.0 .58 0
Alfalfa hay 46.8 .92 0 47.1 1.31 .2
Wheat bran, hard red 48.2 3.29 .2 54.5 3.53 .9
Corn silage 51.5 .10 .2 52.8 1.12 .7
Green peas 15.7 1.12 2.1 16.5 .21 1.1
'Adapted from Jemi et al. (19), values corrected for water content.
2Bucillus subri2is amylase old Sigma Type XIA (Sigma Chemical Co., St. Louis, MO) is used in the ND-S method
described in Table 2.
3R~beWnand Van Soest (%)I; heat stable a-amylase is used in the ND-T method described in Table 2.
4Mean of three replicated experiments.
ethanol or triethylene glycol seems necessary ally, and the mixture is boiled for 1 h and
for concentrate feeds (16). handled as in procedure A.
The NDF method has been criticized for not
recovering pectin, which has beem regarded by Use of Sulfite
some as part of the cell wall matrix. Although
a botanical argument can be made, the evi- The use of sodium sulfite in the NDF pro-
dence from fermentation with gut microorgan- cedure remains optional. Its purpose is to
isms and digestion trials is that pectin is lower the protein level and remove keratina-
unique in being completely and rapidly fer- ceous residues of animal origin. Sulfite cleaves
mentable and therefore is not, in contrast with disulfide bonds and thus dissolves many cross-
hemicellulose, a part of the crosslinked ligni- linked proteins. Its general use for ruminant
fied matrix (45). Pectin also possesses a very feeds is discouraged, especially if the residues
high cation exchange, at least in the de- are to be used as an assay for ND insoluble
methylated form. Our view is that when pectin protein, because the sulfite reaction is nonbio-
deserves recognition, it should be determined logical. The ND and acid detergent (AD) in-
as its own entity. It is, however, a part of the soluble proteins from animal products tend to
nonstarch polysaccharides (NSP)discussed in be indigestible, Sulfite also attacks lignin and
this paper. The method using metu-hydroxy- therefore should not be used in sequential
biphenyl (6)modified by Bucher (7) is specific analyses leading to lignin determination or
for galacturonic acid and is a relatively simple when the residue is to be used for subsequent
procedure that can be conducted along with in vitro digestion with rumen organisms.
other fiber procedures.
Lipid Interferences
RECOMMENDED PROCEDURES
Lipid contents above 10%are a problem for
both N D and AD if a separate oil layer forms
Procedure for NDF in the solution because the detergents are more
Procedure A. A .5-g sample is heated to soluble in the lipid phase than in water. High
boiling in 100 ml of ND plus 50 p1 of heat values of ADF and NDF result. Simple
stable amylase (dietary fiber kit; Sigma cata- removal of lipid may be done by brief heating
logue Number A3306) added before the beaker in ethanol and filtering on the pretared crucible
is placed on heat. Sodium sulfite (.5 g), if to be used subsequently for NDF or ADF.
used, is added at this point. Sample is boiled 1 Contents and crucible are boiled in the NDF or
h and filtered on pretared coarse sintered glass ADF reagent as in the sequential procedure
crucible or Whatman 54 paper (Whatman, (51).
Clifton, NJ).Because of varying soil contami-
nation in forages and feeds, the ash content Filtration
should be reported or excluded from the NDF.
The starch-specific enzyme is stable to boiling, There also are some kinds of samples that
insensitive to EDTA, and approved by the frequently offer filtering problems. These will
AOAC. Samples should be ground through a be minimized if proper filtering techniques are
1-mm screen, but not f i e r , because overgrind- followed. The lowest possible vacuum pressure
ing also can worsen filtration. should be used. Liquid should not be added
Procedure B. An alternate procedure for while vacuum is on. Pressure is released when
removing starch from the most difficult sam- adding liquid. Contents are allowed to settle at
ples is as follows: first the sample is treated least 15 s before admitting vacuum. In this
with 30 ml of 8 M urea plus 50 pl of amylase way, finer matter is fitered onto a settled mat.
added to a 1-g sample; then it is stirred with a The very hottest water is used, and the crucible
rod to break any lumps. The mixture may be should not be allowed to cool. The solution is
heated briefly on a steam bath 80 to 90°C for 5 returned to the beaker and reheated if neces-
min. Then, it is incubated at room temperature sary. If a crucible clogs, positive pressure
for 4 h or overnight and diluted with 100 ml of should be exerted from beneath to flush parti-
ND solution; 50 p1 of enzyme is added option- cles out of the filter plate. This is also a
J o d of Dairy Science Vol. 74. No. 10, 1991
SYMPOSIZTM: CARBOHYDRATE METHODOJBGY 3587
recommended method for cleaning of mi- more economical. Its principal advantage is
bles. Crucibles can be tested after cleaning for that estimates of hemicellulose and cellulose
the speed that liquid will flow. Inherently by difference are more accurate in a sequential
slowly filtering crucibles should be discarded. system. Hemicellulose estimated by subtrac-
tion of ADF from NDF will be too low when
Acid Detergent Flber and Lignin
pectin is precipitated into the ADF. Biogenic
silica has a similar effect because it is soluble
Acid detergent fiber is intended as a prep- in the ND reagents and insoluble in the AD
aration for the determination of cellulose, lig- reagent.
nin, ADIN, acid-insoluble ash (AIA), and sili- Sequential treatment cannot be applied
ca. It is not a valid fiber fraction for nutritional universally because there are specific instances
use or for the prediction of digestibility. Sum- in which fractions of interest can be lost in the
mative systems are mechanistically valid and process. In particular, biogenic silica, AIA,
should replace empirical regressions (8, 45). some tannins, and ADIN are better done on a
'Zhe ADF procedure was collaborated with direct ADF. For tannins, a double sequential
the AOAC (44) and given first action. This analysis can be performed (51) in which ND is
procedure avoided the use of decalin. The Kla- followed by AD and, in parallel, AD followed
son lignin procedure was collaborated at the by ND. Lignin values from these two se-
same time. Since that time, the use of asbestos quences are compared on the two residues.
has been abandoned (36), allowing more flexi- Presence of insoluble tannins is indicated by
ble handling of sequential lignin procedures. higher values from the ND-ADsequence com-
Klason lignin is a better marker than perman- pared with the AD-ND sequence (51).
ganate lignin; however, sequential treatment, Sequential analysis can begin with total die-
i.e., Klason lignin followed by treatment with tary fiber or with NDF as the difference be-
permanganate, yields lignin by difference that tween dietary fiber and NDF as water-soluble
is more recoverable in feces (51). The fraction NSP. The difference should be corrected for
resistant to both 72% (wt/wt) H2SO4 and per- ash and CP (N x 6.25). The crucible contain-
manganate is cutin, which is important in ing the fiber preparation can be analyzed se-
many s e e d hulls and bark quentially using a Tecator (Helsingborg,
Sweden) fiber apparatus or other fiber appara-
Acld-Insoluble Ash
tus using Bemlius beakers. In either case, the
same crucible accompanies the sample
Neutral detergent reagent dissolves pectin throughout the sequence. If Berzelius beakers
and biogenic silica but not silicaceous soil are used, the crucible is placed on its side in
minerals. On the other hand, AD precipitates the beaker, and the sample is boiled in 100 ml
pectic acid as the quaternary detergent salt and of reagent plus enough solution to cover the
quantitatively recovers all silica (51). Acid- crucible. At the end of boiling, the crucible is
insoluble ash is conveniently measured as the removed with tongs, Msed into the beaker,
residue from ADF after ashing at 525'C. It is a placed on the filter, and all liquid is passed
preferable procedure and shorter than that of through the crucible (51).
Van Keulen and Young (42), which is liable to
incomplete recovery of silica due to lack of Total Dietary Flber
sufficient acid dehydration (51). The insoluble
ash after lignin determination by either The concept of total dietary fiber arose as a
KMnO4 or Klason pmcdures is identical to result of interest in fiber and human nutrition.
that of the original ADF, provided that as- It is defined as the polysaccharides and lignin
bestos or other fiiter aids are not used (49). resistant to mammah'an digestive enzymes and
thus is relevant to most monogastric animals
Sequentlal Analysls
with hindgut fermentation. The fractions not
recovered in NDF but resistant to mammalian
The sequential analysis for fiber fractions is enzymes are defied as water-soluble NSP;
attractive because important interferences can they include some legitimate cell wall compo-
be avoided and because the use of sample is nents, such as p-glucans and pectins, as well as
Journal of Dairy Science Vol. 74, No. 10. 1991
3588 VAN SOEST ET AL..
Redpltate
Dry, 1Wc
some storage polysaccharides, such as galac- and Ca salts (from the sample and the buffers
tans in beans and sundry other gums and used), which are insoluble in alcohol, often
mucilages. leads to more ash in the fibrous residue than
The definition of total fiber has led to a p present in the original sample. Volatile loss of
propriate enzymatic procedures isolating the ash components upon ashing at 525'C is prone
fractions resistant to amylases and proteases. to overestimation of fiber.
The first of these is the Asp procedure (35) Because of these problems, various modifi-
adapted and collaborated by the AOAC. This cations of the method have been proposed. We
method digested the heat-gelatjnized sample have developed a new procedure involving
with heat stable amylase, amyloglucosidase, urea-enzymatic dialysis, which avoids heat
and a protease. The final undigested fraction is treatment and removes products via dialysis.
precipitated by 4 vol ethanol. The residue is The principle of the method depends on the
corrected for N x 6.25 and for ash. extraordinary activity of the heat stable en-
The AOAC procedure offers many prob- zyme in 8 M urea. The schematic of the
lems because hydrolyzed products remain in method is shown in Figure 1. Detailed infor-
the solution. Their occlusion by ethanol precip mation on the urea-enzymatic dialysis proce-
itation is a major problem. Interference by Na dure is available (19).
Journal of Dairy Science Vol. 74, No. 10, 1991
SYMPOSIUM: CARBOHYDRATE METHODOLOGY 3589
TABLE 4. A listing of major carbohydrate in NSP' and NSC' (45).
Carbohydrate Mainoccamnce Wata solubility clsssit3~tiOn3
SucrOSe All plants + nonstnrctnral
F~ctans Temperate grass and composites f nonstnrctnral
Slarch Cereal see& - nonstructaral (storage)
Galactans Ltsnme seeds (soybeans) + nonstructud, NSP (storage)
~Glucans Barley OW + sf~clural,NSP
PeClins Lecnrmes and other dicots f structaral. NSP
'Nonstarch polysaccharides.
2Nons~cturaJcarbohydrates.
%om the aspect of function in e plant.
for example, are quite insoluble and even indi- be useful for estimating total m e n balance
gestible (9). In speaking of solubility in refer- and output. The disadvantage is that dietary
ence to physical properties, ease and degree of limits also need to be put on starch and sugars,
solubility of nutrients in the rumen and in because these components are liabilities for
other parts of the digestive tract of farm overproduction of lactic acid (41).
animals have a major effect on dietary quality The general assumption that soluble sub-
and digestive efficiency for both ruminants and stances are more easily and rapidly digested
nonnuninants, although the respective physical than insoluble ones is true only in a general
factors affect these two groups of animals sense. Some insoluble carbohydrates, e.g., un-
somewhat differently. The major factors affect- lignified amorphous cellulose in vegetable
ing solubility and ease of digestion, but not wastes, may be more rapidly fermented than
necessarily intercorrelated, are crystallinity and some of the more soluble modified starches
macromolecular structure. Because these tech- and hemicelluloses.
nical topics transcend the scope of this paper, When large amounts of starch and sugar are
some references are given (4, 9, 12, 39, 45). added, the fermentation pathway can switch to
The soluble NSC are digested rapidly and a lactic acid production, which can lead to
almost completely fermented in the rumen (90 acidosis. However, other soluble NSP, such as
to 100%). The insoluble, resistant starches may pectins, arabans, and Pglucans, are not fer-
escape. It can be argued that an NSC value mented to lactate (41). Hence, there is ment in
including pectin is more appropriate because it distinguishing those feeds containing NSP be-
is a rapidly digested carbohydrate. This would cause these can elicit good rumen efficiencies
18
MATURE CATTLE
AYRSHIRES A
GUERNSEYS G
T
JERSEYS J
Figure 2. hgarithmic plot of rumination capacity (grams) of NDF per minute and BW of cattle. Regression slope for
mature cattle is .95 and not significantly Merent from unity. Regression slope for immatPre and growing animals is
1.50. Data illustrate the greater chewing d c i e n c y of larger animals. Cnldated from data of Bae et al. (5) and Welch
(55) and unpublished figures.
without the problems associated with too much sugar values for many feeds but is larger when
starch (46). The p-glucans that occur in oats the feedstuff contains significant quantities of
and barley contribute to gumminess and are NSP, which includes pectins. Pectins are im-
objectionable in poultry diets (3) but probably portant in citrus, beet pulp, and legume forages
are beneficial in ruminant diets. See Jeraci and but are low in grass forages. Oats, barley, rye,
Lewis (18) for methods for &glucans. and triticale contain bglucans. The NSP can
Because NDF and non-NDF carbohydrates be estimated by difference from total dietary
represent the bulk of most feedstuffs, variation fiber and NDF. The residues should be cor-
in the ratio of non-NDF carbohydrate to NDF rected for protein and ash.
carbohydrate has become a basis for ration
adjustment (22). At least two systems have The Flber Requlmment
been proposed for dairy cattle that utilize
aspects of this concept. One based on soluble Ruminants generally and dairy cattle in par-
carbohydrate has been patented (34), and an- ticular q u i r e adequate coarse insoluble fiber
other based on NDF (31) has been used for for normal rumen function and maintenance of
normal milk fat test. Normal rumen function in
ensuring adequate rumination and efficient
dairy cattle is associated with adequate rumi-
milk production. There is, at present, no sys-
nation and cellulose digestion. These maintain
tem that takes into account the contrasting rumen pH and cellulolytic microorganisms that
qualities of the so-called soluble carbohy- characteristically produce the higher acetate to
drates, the NSC, or the NSP. pmpionate ratios needed for normal lipid me-
when the ratio of forage to concentrate is tabolism in the cow. Daily rumination time is
decreased (less than 50%) in the diet, balano- directly proportional to coarse NDF intake and
ing rations for NSC becomes important for related to body size Figure 2) (5, 55). Other
high producing cows. This problem is more estimations of fiber, e.g., ADF or crude fiber
likely to occur with grass- or corn silage-based are less well related, because only NDF quanti-
diets than with legume hay. Starch and sugars tatively recovers insoluble matrix carbohy-
can be measured directly (22). The net fraction drates, including hemicellulose. The NDF is
can be reasonably calculated by difference us- better related to intake and gastrointestinal fill
ing one of two formulas (34): 1) NSC = 100 - than any other measure of fiber (30, 4 3 , thus,
(NDF + protein + fat = ash) or 2) NSC = 100 - the expectation that the fiber requirement is
[(NDF - NDF protein) + protein + fat + ash]. better expressed in terms of NDF rather than
They have the disadvantage that NSP are ADF or crude fiber. This point is illustrated by
included. They also may not work on silages the experiments in terms of Welch (56) at the
in which sugars are replaced by fermentation University of Vermont, who examined the
products. The second equation recognizes that abiity of various forages to promote rumina-
in some feedstuffs the protein is not totally tion. The best relationship was with intake of
extracted by detergent. The insoluble pmtein in NDF that was correlated at .99 with chewing
NDF is the slowest to be degraded (52) and time.
should therefore be excluded. This calculated The NRC requirements for dairy cattle (33),
NSC is quite close to determined starch and while allowing different levels of fiber relative
J o d of Dairy Science Vol. 74, No. 10, 1991
3592 VAN SOEST ET AL..
to production, set fixed levels of NDF relative TABLE 6. Corntation coefficimts (r) of cation exchaqe
to ADF for lactating cows. This system fails to capacity detemhed with coppa 0 and praseodyrm'um
0 at specifed p H d batch-isolated neutral-detergent
reflect that the ratios of hemicellulose to cellu- fibers (26).
lose vary widely between feed fiber sources,
particularly in the all important corn silage and Cum R O
alfalfa that are the basis of many rations ('"able Parameter OH 3.5 DH 3.5 ~~~
OH 7.0
5). The recommendations are inconsistent with Li- 96 .76** .69** &a**
wellestablished knowledge. It is apparent that Hemicellulose, 96 .49 .56t .48
the NRC levels for ADF were based on alfalfa, Celldose, 5% .ll .03 .16
because they do not fit Corn silage. Use of the N, 5% .70* sot .5St
Cu at pH 3.5 .%** .95**
NRC recommendation for ADF will result in R at pH 3.5 .we*
overfeeding of fiber in the case of corn silage,
grass silages, and hays. tP < .lo.
Although coarse NDF of any sort will *P < .05.
satisfy the ruminarion requirement, the quality **P < .01.
of that fiber has important effects on the rumen
environment and on microbial efficiency. Be-
yond particle size and adequate levels of NDF,
additional factors of buffering capacity, cation bly magnesium). Cation exchange is the ability
exchange, and fermentation rate are important of fiber to bind metal ions on its surface in
feed and fiber properties needing considera- much the same way that clay minerals are able
tion. Because NDF is not a uniform material, to hold cations in soil The exchange waves as
other physicochemical descriptions become a bank, exchanging K+,Ca*, Na+, and Mg*
important nutritional considerations. The qual- for H+ when pH drops and recharging when
ity of NSC and the proportion of starch and new cations become available as saliva and
sugar relative to other NSP, such as pectin, ingesta are mixed. An advantage of this
have major influences on rumen microbes and regenerative bank is that ruminated fiber, as it
efficiency. These factors, net fermentation rate, passes down the digestive tract, contributes
type and amount of fibrous and nonfibrous buffering action farther down the gut. If the pH
carbohydrates, along with the N and protein rises in the m e n , the bank is recharged with
supply interact to affect rumen function and metal ions, and the bound ions are prevented
microbial efficiency. from washing out of the m e n by their attach-
ment to c o m e fiber.
Mature legume forages are the most effec-
Cation Exchange and Buffering tive dietary ingredients for supplying ex-
The most important feed characteristic be- changeable buffering capacity (Table 7), al-
sides particle size that contributes to net rumen though some concentrate fibers are as good or
buffering is the buffering capacity of the feed; better. Corn silage has only about onethird the
this depends on cation exchange capacity of
capacity of alfalfa and also contains starch,
the fiber and, to some extent, on the fermenta- which can promote lactic acid production.
The buffering capacity of feedstuffs derives
tion of protein to ammonia. Ionexchangeable in part from the physical effects that they elicit
groups in plant cell walls include carboxyl, in the m e n and on rumination. Because the
amino, free aliphatic hydroxyls, and phenolic fermentation of carbohydrates leads inevitably
hydroxyls, all of which have some affinity for to production of large amounts of VFA, their
binding of metal ions ('"able 6). Thus, the removal by absorption and the recycling of
surface properties of fiber, hydration, and ca- mineral ions are essential processes in the
tion exchange are intercomelated (r = -.7) and maintenance of pH and nOnnal rumen environ-
are likely associated with short lag times and ment. Fiber is among the more slowly digest-
rapid rates of cell wall fermentation. Microbes ing solid fractions and contributes most to the
that have negatively charged cell walls (23,24) maintenance of normal rumen environment.
"recognize" fibrous particles through their ex- More rapidly fermenting feeds yield organic
changeable surface and form attachments (l), acids at a faster rate, thus taxing the buffering
which require divalent cation liganding (proba- system to a greater degree. Mature grasses are
Journal of Dairy Science Vol. 74. No. 10, 1991
S Y M P O S I U M : CARBOHYDRATE METHODOLOGY 3593
TABLE 7. Cation exchange capacity (CEC) values for a poor in exchange and buffering capacities but
range of foodstuffs and intakes of NDF and DM required also ferment more slowly. Thus, supplement-
to yield equivalent exchange capacity of 1 mol (100 g)
calcium carbonate (27).
ing grasses with starchy concentrate supple-
~ ments that ferment faster and can yield lactic
Calcium acid renders the rumen more susceptible to
carbonate acidotic conditions that limit rumen efficiency
equivalent
and net feed intake. Under these conditions.
NDF DM
Feedstuff NDF CEC Basis Basis
grass-based forages are less efficient.
There are several systems for measuring
(46) (mes/loog) --I- cation exchange in plant cell walls. Direct
Alfalfa hay 45 50 4 9 measurement of H+ exchange with acid is apt
Birdsfoot trefoil 65 30 6 10 to degrade sensitive carbohydrate structures
coastal bermuda
grass 70 11 17 25 and produce artifacts, although direct titration
Corn silage 44 15 13 30 gives good information (Figure 3). Our first
Cottonseed meal 29 57 4 12 values involved the use of lithium binding
Distillers grabs 50 35 6 11 (50). Lithium is weakly bound, and variability
Dried brewers
Brains 62 29 7 11 is encountered in the washing procedures to
Guineagrass 72 22 10 13 remove unbound lithium. Calcium and barium
Haycrop silage 43 25 8 19 proved unsatisfactory because of sulfate inter-
Oats 37 17 12 31 ference. Later values were obtained with Cu*
Rapemximeal 26 100 2 8 (28) via a modification of the procedure of
Reedcana~ygrass 49 21 4 12
RYeeTm 41 24 8 20 Keijbets and F'ilnik (20). This method is lim-
Safnower meal 60 20 10 16 ited to measurement at pH 3.5 because of the
Soybean meal 12 41 5 40 instability of Cu* at higher pH. More recent-
Sugarbeetpulp 51 70 3 5 ly, the stronger binding rare earth ions (praseo-
Sunflowermeal 19 37 5 29
Timothy hay 63 30 7 11 dymium and neodymium) have been applied in
m e a t -st& 80 13 15 19 a new procedure (2). Values for cation ex-
7.0
6.0
k0
5.0
Q
4.0
2.0
I 1 I I 1 1
0 io 20 30 40 50 60 7Q 80 9G I00
(5,
Y
\ 500
0-
Q,
E
v
z
0
U
U
U
CY 250
W
a
4
0
0
LI / ’
2 ATP
I
PECTIN
1 ---)
I
STARCH 1 PROCESSED
*
- ’1 0
I
CELLULOSE
I
10
I
20
1
30 40
I I
50
RATE O F FERMENTATION, %/h
Figure 5. Tbe relationship between the amount of microbial protein produced per unit of feed fermented in the f11113en
in relation to rate of fermentation. Bacteria in a nonnal nunen ferment carbohydrate to W A with a yield of 4 ATP from
1 glucose. Lactic acid production (lower curve) is characteristic of acidic m e n s and yields only 2 ATPhnol of glucose.
nance cost. The rate of digestion dilutes the degrading starches appear to inhibit cellulose
maintenance cost for aIl rumen bacteria, and digestion, but pectins may impose this penalty.
more rapidly fermenting carbohydrates im- Pectin is high in citrus, beet pulp, and alfalfa,
prove rumen efficiency. When large amounts but there is very little pectin in most grasses
of starch are added to the diet, digestion rates and corn silage.
in rumen fluid increase, and starch digesting The character of pectin fermentation results
organisms like Streptococcus bovis can switch not only from a lack of lactic acid output but
from acetate production, when they get about 4 also from the nature of the galacturonic acid
ATP per unit of glucose fermented, to lactate structure that provides potential buffering
production when they get only 2 ATP per unit through cation exchange and metal ion bind-
of glucose. In this case, the microorganisms ing. Alfalfa contains 5 to 10% pectin as cal-
tend to sacrifice the efficiency of ATP produc- cium pectate, and larger amounts occur in cit-
tion for the sake of increased lactic acid, which IUS and beet pulp as the methyl ester, which is
makes the environment more favorable for hydrolyzed in the rumen to produce metal ion-
their exclusive growth. Streprococcus bovis binding capacity. In view of the faster fer-
produces lactic acid when the pH is low, espe- mentation rates of pectins, these physicochemi-
cially if the dilution rate will be slow. At low cal characteristics probably account for some
pH, growth rates of all organisms decrease, but of the “magic” effect observed when pectin-
cellulolytics are more adversely affected (47). containing feeds are added to high starch diets.
Pectin invariably is the most rapidly Cereal grains are the basis for much animal
degraded complex carbohydrate, whereas feeding, and, because starch is the major com-
starches and celldoses are quite variable ac- ponent, starch quality affects feed efficiency.
cording to source; hence, their quality reflects Starches vary in seeds according to phys-
digestion rate. Selecting different carbohydrate icochemical structure. Linear forms such as
sources to be complementary may be benefi- amylose are more crystalline and are digested
cial, provided that competition among sub- more slowly. There is much genetic variation
strates is not severe. Sugars and rapidly in cereal grains, which may account for differ-
Journal of Dairy Science Vol. 74, No. 10, 1991
3596 VAN SOEST ET AL.
ences between sources. For ease of degradation 6Blumencrant2, N., and G. Asboe-Hansen. 1973. A
new method for quantitative determination of uronic
(43,uncooked starches rank in the following acids. Anal. Biochem. 54484.
order: wheat, barley, oats greater than corn, 7 Bucher, A. C. 1984. A Comparison of solvent systems
and sorghum greater than legume. Response to for extraction of pectic substances from fruits and
processing is in approximately the reverse or- vegetables. M.S. Thesis, Cornell Univ., Ithaca, NY.
8 Conrad, H. R.,W. P. Weiss, W. 0. Odwongo, and W.
der (15). The potential rate of fermentation of L. Shockey. 1984. Estimating net energy lactation
all carbohydrates largely determines their fate from components of cell solubles and cell walls. J.
in the digestive tract and the efficiency with Dairy Sci. 67427.
which microbes can use them. 9Englyst, H. N., and G. T. McFarlane. 1986. Break-
down of resistant and readily digestible starch by
human gut bacteria. J. Sci. Food Agric. 37:699.
CONCLUSIONS lOFonnesbeck, P. V. 1976. Estimating nutritive value
from chemical analyses. Page 219 in 1st Int. Symp.
Fiber has come to be recognized as a re- Feed Composition, Animal nutrient requirements and
quired dietary ingredient for many herbivorous computaization of diets. P. V. Fonnesbeck, L. E.
Harris. and L. C. Kearl, ed. Utah State Univ., Logan.
animal species and is necessary for normal 11 Fox, D. G., C. J. Sniffen, J. D. O’Connor, J. B.
rumen function in ruminants. Quality of fiber Russell, and P. J. Van Soest 1990. The Cornell net
varies according to fermentability, particle carbohydrate and protein system or evaluating cattle
size, and buffering capacity. Only coarse in- diets. Search Agric. Cornell Univ. Agric. Exp. Stn.
No. 34, Ithaca, NY.
soluble fiber is adequate for promoting rumen 12 French,D. 1973. Chemical and physical properties of
function. This corresponds to the NDF from starch. J. Anim. Sci. 37:1048.
forages, and NDF is the preferred measure for 13 Giger. S., M.Dorlutns, and D. Sauvant. 1981. Adag
ruminant feeds and dietary balancing pro- tation of the Van Soest method to a routine determina-
tion of concentrate feedstuffs. Comm. Eur. Commuu
grams. Therefore, the standardization of proce- Workshop Methodol. Anal. Feedingstuffs Ruminants.
dures for NDF is of paramount importance. European Van Soest Ring Test. Rep. meeting to dis-
Recommended procedures have been provided cuss analytical results. Ministry of Agriculture, Fisher-
Nonstructural carbohydrate in ruminant ies, and Food, Slough Laboratory, Lond., Engl.
14Goering, H.K.. and P. J. Van Soest. 1970. Forage
feeds also can have impact on dietary quality fiber analyses (apparatus, reagents, procedures, and
and microbial efficiency in the rumen, and some applications). Agric. Handbook No. 379. ARS-
computerized systems for using both NDF and USDA, Washington, DC.
the NSC have been proposed (11, 34). The 15Hale. W. H. 1973. Influence of processing on the
utilivltion of grains (starch) by ruminants. J. Anim.
NSC can be further subdivided into those car- Sci. 37:1075.
bohydrates (starch and sugar) capable of yield- 16Jeraci J. L., and P. J. Van Soest. 1990. Improved
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acid, because lactic acid production has major fiber. Adv. Exp. Med. Biol. 270245.
17 Jeraci, J. L., T.H. Hernandez, J. B. Robertson, and P.
impact on rumen efficiencies. The latter (NSP) J. Van Soest. 1988. New and improved procedure for
include pectins, galactans, and p-glucans. neutral-detergat fiber. J. Anim. Sci. 66(Suppl. 1):
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18 Jeraci, J. L.,and B. A. Lewis. 1989. Determination of
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