Cellulases and Their Applications

18 The Value of Wood-Derived Products in Ruminant Nutrition

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W I L L I A M H . P F A N D E R , S. E . G R E B I N G , G E O R G E HAJNY, and W E N D E L L TYREE Missouri Agricultural Experiment Station, Columbia, Mo. and Forest Products Laboratory, Madison, Wis.

Literature reviewed indicated that raw wood, as browse, and products such as leaves and beans can be used in ruminant rations. Raw woods are poorly utilized as energy sources. Cellulose prepared from wood is digestible, but particle size limits utility. Wood molasses and other hydrolytic products are well digested. Hemicellulose (HC) was obtained from mixed hard and soft woods. The distribution of the sugars was: galactose, 7.1; glucose, 14.4; mannose, 26.0; arabinose, 4.9; and xylose, 47.4. The HC was also neutralized and/or dried to 95% dry matter. All products tested at 30% of the ration were non-toxic to wethers for five years and to ewes through two reproductive cycles. The estimated digestible energy, in kcal./kg., for the products was: 65% DM from Laurel, 2900; 65% DM from Ukiah, 2420; 95% DM, 3100.

hen an animal scientist looks at wood, he can elect to view it i n " several ways: (a) as an indigestible, highly abrasive material that increases fecal dry matter and nitrogen excretion, ( b ) as a mixture of cellulose and lignin from which, if, the lignin envelope could be removed, a ruminant could obtain a digestible cellulose, and ( c ) as a complex mixture consisting of ( b ) plus a rich supply of hemicellulose and a multitude of other compounds. In classical and current literature, there is support for each view. W e w i l l consider each i n turn and then present in some detail the estimates of nutritional value and limitations of some derived products. 298

In Cellulases and Their Applications; Hajny, G., et al.; Advances in Chemistry; American Chemical Society: Washington, DC, 1969.

18.

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299

Ruminant Nutrition


The bases for most ruminant rationing systems are rooted i n the German schools of the 1800s. Variations of the Weende analytical scheme are included i n A O A C (2) procedures and variations of the net energy scheme of Kellner (33) are used as the "ultimate" measure of energy yield of ruminant rations. Both systems have limitations and have survived numerous critiques. W o o d is a complex product of an unstable biological system. Its total chemical composition and the physical-chemical relationships remain to be elucidated. Parts of the tree are normally eaten at some seasons of the year by ruminants and other herbivores. Bark, browse, and leaves are the major items consumed (6, 19, 27, 28, 29). The reported composition of several components are shown i n Table I. Table I.

The Composition and Nutritive Value of Wood Products DM %

Beech leaves Birch leaves Elm leaves Oak leaves, dried Poplar leaves Brush wood Brush wood, beech Acorns Locust beans Sawdust from pine Molasses, wood Molasses, cane° Red clover hay*

43 45 88.0 94.0 84 75 84.7 85.0 85.0 83.5 60.0 74.0 83.5

NFE %

DP %

Protein COD*

9.8 6.9 8.6 29.9 17.4 26.7

21.7 24.7 49.9 45.3 39.6 40.3

6

11.6 6.4 62.4 0 0 22.2

61.6 69.0 19.6 55.6 62.0 35.8

3.4 3.9 8.5 0 3.4 1.6 0.1 3.8 3.2

Protein Fiber % % 6.9 7.9 15.9 9.3 10.8 4.6 4.0 5.7 5.8 0.3 0.5 3.0 15.3

0 0 7.0

73 56

SE TDN* TDN e

d

19.2 26.0 50.0 26.7 14.5 -12.9 69.0 71.7 -3.3 35.6

17

49 54 54

° Common feeds for comparison. A reported value considerably different from that used to calculate the DP figure. Ref. 33. Adapted from Ref. 40. b e

d

Kellners investigations which showed that fiber content limits the net energy value and that added sawdust depresses utilization of other components of the ration have discouraged research opportunities. Perhaps the most damaging data are those leading to the conclusion that the addition of 100 grams of sawdust had the same effect as removing 3.2 grams of starch (33). A shortage of hays and concentrates i n W o r l d Wars I and II stimulated research into the possibilities of preparing hydrolyzed sawdust or "fodder cellulose" from wood. Some reports showed good utilization of these products, but most of the work indicated that the value of the



300

CELLULASES A N D THEIR APPLICATIONS

derived products was closer to that of sawdust than of hays (3, 8, 22, 23, 25,31, 38,44,47,49,50,51). Generally, the rations used were not fortified with a full complement of required minerals, and some disturbances developed. More recently, sawdust has been used to limit feed intake or to serve as a source of "roughage" i n the ration. T e n to 15% of sawdust can be fed without evidence of toxicity (16, 45). Nearly pure cellulose derived from wood has been used extensively i n semi-purified diets (20). The digestibility of the product is limited i n ruminants, but under in vitro conditions it is nearly 100% digestible if the reactions are continued for long periods (36). The availability of this product for microbial fermentation suggests that if an economical way of removing lignin were available, considerable wood would find its way into ruminant feeds. In fact, newspapers are now being used i n experimental feeds (34). Prior to and during W o r l d W a r II, research was initiated to develop a source of wood molasses for alcohol production. Later, these products of acid hydrolysis were fed to ruminants with good results (5, 14, 15, 17, 24,32,35,40,41). Table II.

pH Ash Solids Sugars (as is) Sugars (after hydrolysis) Methoxyl Lignin Acetyl Uronic Anhydride % of sugars As is Galactose Glucose Mannose Arabinose Xylose After hydrolysis Galactose Glucose Mannose Arabinose Xylose

Composition of Hemicellulose

L 4%

L 43%

3.80 0.15 4.00 0.79 2.35

4.04 1.52 8.09 25.56

0.35

2.00

16.03 6.34 10.71 45.09 21.83

17.58 10.88 11.43 31.95 28.16

6.58 15.54 27.92 5.53 44.43

7.84 14.19 27.33 5.11 45.53

LHC 1 2.17 64.00 34.14

LHC 2 4.08 2.17 64.00 12.27 37.24

8.50 3.75

7.85 14.34 27.17 4.88 45.76

6.37 14.57 24.82 5.08 49.16

°The following abbreviations are used: L—Laurel, Mississippi, Raw material about softwood; HC—-Hemicellulose; N—Neutralized.



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Ruminant Nutrition

The main direction of research now appears to be toward the utilization of wood by-products. W o o d generally contains from 50-60% cellulose, 20-30% lignin, 10-20% pentosans, 0 - 7 % mannans and some acetyl linkages (usually 0.5-2% acetic acid) (48). This wide assortment of carbohydrates is of interest to ruminant nutritionists for at least three reasons. Ruminants need a source of readily available energy, usually obtained as hexoses derived from soluble carbohydrates. Most forages contain pentosans or hemicelluloses, but no reliable concentrated pure source of these materials is available and until recently it was not possible to obtain enough to evaluate the utility of these products i n defined rations. Finally, the possibility of producing polymers from feed material is attractive. Studies designed to determine the role of some of these by-products in ruminant rations are described i n the sections which follow. Methods and

Experimental

The raw material was obtained as a by-product from the manufacture of hard board. In the usual process, logs are debarked and fed into a Concentrates' at Several Production Stages NLHC 1 7.00 7.40 63.70 9.78 31.03 1.92 5.88 7.09 4.15

NLHC 2

65.00 11.64 34.98

Dried UHC

Dried LHC

NUHC

4.76 97.10 18.72 58.40 3.06 5.35 7.36 6.86

0.86 64.00 24.50 42.48 1.14 0.92 4.26 2.32

18.08 56.26 2.41 7.44 9.13 5.76

5.98 16.40 15.96 3.99 57.67

13.31 17.11 44.66 3.96 20.95

6.67 15.84 25.75 5.56 46.18

4.96

9.66 13.17 12.55 29.49 35.13 6.71 16.09 24.06 4.13 49.01

5.81 14.97 25.16 4.91 49.15

60% mixed hardwood and 40% pine; U—Ukiah, California, Raw material about 90%


302


chipper. The chips are subjected to live steam at 42 kg./cm. pressure for one minute and washed i n counter current washers. The washings contain about four percent dry matter. The washings are evaporated to 40% solids and then to 65%. The p H of the 65% solids product is about 3.7. It is referred to as hemicellulose ( H C ) (trade mark, Masonex, Masonite Corporation, Chicago, Illinois). If desired, the material may be neutralized ( N ) to p H 5.5 or spray dried. The typical raw material at the Laurel, Mississippi ( L ) plant contains 60% mixed hardwood and 40% pine. The Ukiah, California ( U ) plant processes about 90% softwood. Table II shows some typical compositions obtained by modifications of the methods of Saeman et al. (46) from products produced i n various ways. Results of typical feed analyses are shown i n Table III. Palatability Trials. T w o to five sheep were placed i n a pen containing a feeder with four movable compartments. The compartments were numbered and their positions randomized each day. T w o feeds were compared i n each pen. Each day a known amount of feed was placed i n each compartment and the position was determined at random. Twenty-four hours later the refusals were removed, weighed, and fresh feed added. A normal test period was seven days. The first trial was designed to detennine the relative palatability of hemicellulose and molasses. The basal ration consisted of timothy hay, 68%; corn, 28.16%; 44% C P soybean oil meal, 2.31%; urea, 0.77%; dicalcuim phosphate, 0.26%; salt, iodized, 0.50%; and vitamin A and D concentrate to meet N R C requirements. Other rations were prepared by removing corn and increasing the soybean meal and adding 5 % dry matter from cane molasses, L H C , dried molasses, or dried L H C .


2

Table III.

Typical Results of H C Analyses According to Current Feed Control Methods

Component Protein Fat Fiber Ash Solids Calcium Phosphorus

HC .5% .5% 1.0% 6.0% 65.0% .50 .07

Dried HC .5% .5% 1.0% 4.0% 97.0% .50 .07

The basal ration i n Trials 2 and 3 was the same as i n Trial 1 and the test materials were added at 5 % of the ration dry matter with corresponding adjustments i n corn and soybean meal to maintain isonitrogenous diets. In Vitro Studies. A series of artificial rumen trials was run to test the effect of neutralization and hemicellulose source on normal cellulose digestion. The procedure used was that of Hargus (21). Feeding Trials. T R I A L 1. Six lots of nine lambs each were used to compare hemicellulose and cane molasses. Each treatment was replicated three times. The rations used contained 66% ground ear corn, 16.6% mixed hay, 10.4% soybean oil meal, and 7% of either cane molasses or



18.

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Ruminant Nutrition

H C . A mineral mixture was available. Gain, feed efficiency and carcass characteristics were determined. T R I A L 2. T w o lots of eight lambs each were used to determine the value of dried H C as a roughage. The basal ration contained 10% cottonseed hulls, corn, urea, minerals, Aureomycin and stilbestrol. The ration was calculated to supply 12.5% protein, 0.4% calcium, 0.3% phosphorus, 0.6% potassium, and 3.0% fat. I n the test ration, 8 % dried hemicellulose and 0.15% urea were substituted for 5 % hulls and 3.15% corn. About 100 pounds of long hay per lot was supplied during the first week the lambs were on test. From that time on, they were self fed their rations i n sand covered lots that had automatic heated water cups. They were killed after 66 days and hot carcass weights were obtained. Twenty-four hours later, cold carcass weights and grades were obtained. T R I A L 3. Twelve percent of dried L H C replaced ten percent of cottonseed hulls i n a fattening ration for lambs. The remaining ingredients were similar to those used i n Trial 2. Evaluation of Energy Availability. T R I A L 1. The rations shown i n Table I V were designed to estimate the value of hemicellulose i n rations for lambs and mature wethers. Each sheep received each ration for about 30 days. During the last 14 days of each period, animals were i n digestionmetabolism stalls to determine nitrogen and energy use. At the completion of each series the acceptability of rations was measured by the time required to change to the new treatment. T R I A L 2. A 2 X 2 X 2 factorial design was used. The variables were hemicellulose from Laurel, neutralized or unneutralized and hemicellulose from Ukiah, neutralized or unneutralized, each fed at 15 and 3 0 % of the ration. The rations are shown i n Table V . There were four sheep per treatment. The standard procedure for digestibility trials as developed by the informal forage evaluation committee (30) was used. Each animal was fed individually on the ration for approximately three weeks during which time an estimate of the voluntary feed consumption was made and during days 21 to 30, a standard digestibility trial was completed. Table I V . Composition of Rations Used to Evaluate Digestibility of H C , T r i a l 1

Ration Timothy hay Shelled corn, ground Soybean meal, 44% Urea Dicalcium phosphate Ground limestone Salt Molasses Hemicellulose Vitamin A, 5000 IU/gm. TOTAL

Basal % 67.78 28.95 1.81 .77 .11 .08 .50

— — 20 gms.

100.0

Basal + Molasses %

Basal + HC % 61.01 26.07 1.61 .69 .10 .07 .45

61.01 26.07 1.61 .69 .10 .07 .45 10.0

—

10.0 20 gms.

—

20 gms. 100.0

100.0


304


A t the conclusion of the experiments, the animals were changed immediately to the next ration to be tested. T R I A L 3. A t the conclusion of the factorial design, four sheep received 50% Laurel hemicellulose and another group of four sheep received i n sequence the high energy ration and then the high energy ration supplemented with 15 and 3 0 % dried hemicellulose. The composition or the rations is shown i n Table V I . Composition of the rations was determined by standard A O A C procedures ana the energy content by methods previously reported b y mis station (21).


Table V . The Composition of Rations Used to Evaluate the Energy Value of Rations in Trial 2 Level of Hemicellulose Ingredient, %

0

15

30

50

Cottonseed hulls Hemicellulose

85.3 0

70.0 15.0

55.3 30.0

35.3 50.0

The rations all contained: soybean meal, 13%; dicalcium phosphate, 0.7%; salt, iodized, cobaltized, 0.5%; Vitamin A and D premix, 0.3%; sodium propionate (as a mold inhibitor), 0.2%. Table VI. Composition of Rations Used to Evaluate Dried Hemicellulose Level of Dried Hemicellulose Ingredient, Kg

0

15

30

Corn, ground Urea, 42% N Hemicellulose, dried

955.00 13.80 —

809.00 17.73 150.00

655.00 21.63 300.00

Tests to Determine Long Term Effects. I n 1963 wethers were placed on rations containing 3 0 % H C . They have been maintained i n confinement and have received no other foods. Sixteen yearling ewes were allocated into two equal groups and fed the rations shown i n Table V I I . They were mated to Hampshire rams i n September of each year and carried through two gestation-lactation periods i n drylot. Observations were made on the general performance of the ewe and on the vigor and growth rate of the lambs produced. During the second year, lambs were killed and detailed tissue samples were obtained for further evaluation. Electrophoretic patterns on the ewe's blood and milk were obtained.

Results Palatability. Results are shown i n Table V I I I . As the sheep i n creased i n size, their feed intakes also increased. The various lots of sheep were not always consistent in their preference. Lot D seemed most sensitive and G least sensitive to ration variables. Some of each ration



18.

305

Ruminant Nutrition

PFANDER E T A L .

was always consumed and even the relatively unpalatable ration (4) was consumed in adequate amounts when the animals had no other choice. Sensitivity may have decreased as the experiment progressed. The sheep in these trials preferred hemicellulose and dry molasses. In two groups, D and G , there was a marked preference for molasses instead of the basal ration. N o group expressed a strong preference for hemicellulose as compared to the preference for the basal ration. However, wet hemicellulose was well accepted by all lots in all comparisons. The results of Trial 2 are shown in Table IX. In this test, the unneutralized product was more palatable than the neutralized and the basal ration was preferred to either product. Results of Trial 3 are shown i n Table X . If only overall averages are examined, the lambs appear to prefer the unneutralized products, but this is not a true indication of the values obtained. In the first place, the intake of these lambs declined as the weather became warm in July and August, and the unneutralized products were fed early i n the trial. W h e n the dry products were compared, the sheep d i d prefer the unneutralized product; but, of the wet products, they preferred the neutralized material. In Vitro Trials. There were no significant effects on cellulose digestion where H C was added at levels of 1 - 1 5 % to the standard assay system. Feeding Trials. T R I A L 1. The data are summarized in Table X I . There were no differences between the ration containing H C and that containing molasses. T R I A L 2. The results are shown in Table X I I . The performance of the dried H C lot is equal or superior to the control. Table V I I .

Composition of Rations Used for Reproduction Studies Ration Ingredients, Kg

Alfalfa, meal, reground pellets Cottonseed hulls Ground corn Soybean meal Cane molasses L H C , 65% solids Dicalcium phosphate Iodized, cobaltized salt Limestone Sodium sulfate Vitamin A & D—2250, 400 IU/gm. Urea, 262 plus 0

TOTAL

I

2

100 500 208.5 90 75

100 500 206.5 90

— 5

— 75

5 2 0.5 1.0 13.0

5 5 2 2.5 1.0 13.0

1000.0

1000.0

• Contains 45% N, product of E. I. du Pont.


306


Table VIII.

The Palatability of Rations Containing Comparison

Rations 2 vs. 3 Groups

Rations 3 vs. 5

Rations J vs. 2

kg.

kg.

kg.

kg.

kg.

kg.

kg.

kg.

15.2 14.1 16.8 10.3 16.0

10.4 9.4 14.0 14.5 8.7

2.5 7.1 14.9 5.0 8.7

25.1 22.8 19.6 23.2 19.6

19.1 13.0 19.1 17.7 13.7

13.9 19.3 21.5 17.0 17.4

13.5 15.2 15.7 14.3 14.3

18.4 13.0 20.7 15.0 15.2

Sum

72.4

57.0

38.2

110.3

82.6

89.1

73.0

82.3

Mean/group

14.5

11.4

7.6

22.1

16.5

17.8

14.6

16.4

0.7

0.5

0.4

0.8

0.8

0.7

0.8

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Rations 4 vs. 5

Mean daily/head Ration No. Ration No. Ration No. Ration No.

1= 2= 3= 4=

1.0*

Basal Basal plus dried cane molasses Basal plus cane molasses Basal plus dried hemicellulose

Table IX.

Intake of Feeds With and Without Hemicellulose Rations

Lot

NL vs. UNL

Basal vs. NL

Basal vs. UNL

D E G H I

19.7 18.7 15.6 17.3 9.0

19.5 21.4 15.2 20.5 10.4

21.1 20.6 23.7 23.5 27.0

15.6 9.4 9.3 15.2 8.5

25.2 19.2 19.4 23.8 15.6

11.0 19.0 21.2 14.1 10.4

Sum

80.3

87.0

115.9

58.0

103.2

75.7

Mean*

16.0

17.4

23.1

11.6

20.6

15.1

In kg/lot/week. NL = Neutralized Laurel UNL = Unneutralized Laurel

a

T R I A L 3. The results are shown in Table XIII. The dried H C has apparently replaced all "roughage" i n the lamb fattening ration. In both direct comparisons, the lambs fed hemicellulose gained faster and more efficiently than those fed cottonseed hulls. Representative carcasses are shown in Figure 1. It is obvious that the hemicellulose fed lambs are equal or superior to the controls. Evaluation of Energy Availability. T R I A L 1. The results are shown in Table X I V . N o differences are significant (P > .05). The slightly


18.

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Ruminant Nutrition

Five Percent Dry Matter from Molasses or Hemicellulose Comparison


Rations 3 vs. 4

Rations 1 vs. 5

Rations 4 vs. 4

Rations 2 vs. 5

Rations 1 vs. 4

kg.

kg.

kg.

kg.

kg.

kg.

kg.

kg.

kg.

kg.

30.6 13.5 15.0 17.8 14.4

3.7 17.1 21.5 15.7 13.6

22.0 15.8 23.2 19.1 16.7

17.3 17.7 22.2 16.4 16.8

16.2 15.8 18.5 16.6 14.8

17.1 16.4 20.5 17.2 16.6

21.3 22.0 24.5 23.0 19.1

23.8 14.3 18.8 14.8 15.0

20.3 22.7 12.8 18.6 27.8

12.9 20.4 18.6 18.9 10.0

91.3

71.6

96.8

90.4

81.9

87.8

108.9

86.7

102.2

80.8

18.3

14.3

19.4

18.1

16.4

17.6

22.0

17.3

20.4

16.2

0.9

0.7

0.9

0.9

0.8

0.8

0.8

0.9

0.8

1.0*

Ration No. 5 = Basal plus hemicellulose • Significant difference between pairs, student's "t" < 0.01. Significant difference between pairs, student's "t" < 0.05. 6

lower protein digestibility of the molasses and H C rations is characteristic of such rations supplemented with carbohydrates. T R I A L S 2 A N D 3. The results of Trials 2 and 3 are shown i n Table X V , and the analysis of variance of the individual data are reported i n Table X V I . There was a highly significant difference i n level and source of hemicellulose, but no effects from neutralization. In Trial 3, there was a marked depression of digestibility at the 30% level of dried hemicellulose. It was calculated by the difference procedure that the digestible energy of the Laurel hemicellulose is 2900 kcal./kg., and of the Ukiah hemicellulose 2420 kcal./kg. A n interaction between source and level complicates the interpretation of these data. D r i e d hemicellulose when fed at 15% of a high energy ration is estimated to provide 3100 kcal./kg., but the 30% level depressed digestibility and reduced the digestible energy content of the ration. Long Term Effects of Feeding H C . The wethers have completed five years on their rations. N o losses have occurred. There were no differences in the reproductive performance of the two groups of ewes and no unusual disturbances were noted i n any situation.

Discussion The literature reviewed indicates that while wood is regarded as a possible component in emergency rations, it is not highly regarded for animal production. However, it has been stated that, "probably more


308


Table X. July 2-7 Dry vs. Wet HC HC


Group

Total

Intake of Lambs Fed Laurel July 27-Aug. 2 Wet vs. Wet HC NHC

Total

G E I H

1631 1403 1743 1735

1483 1416 1430 1855

1393 1226 1258 1438

1568 1304 1912 1798

Sum

6512

6184

5315

6582

Mean'

1628

1546

1329

1645

Total, all dry Total, all wet Total, all neutralized Table XI.

3174

2974

5460 6016 5492

Result of Feedlot Performance According to Kind of Ration Molasses

No. of lambs Av. initial wt. (kg.) Av. finished wt. (kg.) Days on Feed Av. daily gain (kg.) FeedAg- gain Av. carcass grade Av. carcass yield, %

27 35.49 47.01 63 .18 4.09 9.4 C + 52.88

Hemicellulose 27 35.48 47.01 63 .18 4.11 9.2 C + 52.83

Table XII. Lamb Performance when Dried Hemicellulose Replaces Part of the Cottonseed Hulls in a Lamb Finishing Ration

No. lambs Starting weight, av. Finish weight, av. St. Louis weight, av. Carcass weight, av. Ave. daily gain Percent shrink Yield, percent Grade Condemned livers Feed efficiency b

6

d

6

c

Control

Dried LHC

8 30.700 kg. 48.900 kg. 47.900 kg. 25.600 kg. .275 1.740 53.460 8 4 5.150

8 31.400 kg. 52.200 kg. 50.200 kg. 26.900 kg. .311 3.650 53.550 8 3 5.320

* Spray dried. * Weighed after feeding or full weight. ° Weighed after 18 hours without feed. Used to calculate percent shrink to market and yield. Cold carcasses about 22 hours after killing. Used to calculate yield. * "Saw dust" and/or tapeworms in controls. Tapeworms in HC lot. d


18.

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Ruminant Nutrition

309

Hemicellulose Processed in Different Ways


Aug. 16-21 Dry vs. Dry HC NHC

Total

Aug. 22-27 Dry vs. Wet NHC NHC

1468 1418 1693 1343

1093 1193 1206 1243

1212 963 1433 1062

1408 1374 1408 1792

5922

4735

4670

5982

1481

1183

1168

1496

2664

Total, all unneutralized 5984 • Each value is average of 3 lambs per lot. Table XIII. The Value of Substituting Dried H C for Cottonseed Hulls in High Grain Rations Lot # 10 11 9 7

Variable

ADG

Feed Efficiency

HC Control HC Control

.190 .185 .212 .178

6.6 7.6 6.1 7.8

Figure 1. Representative carcasses from lambs fed hemicellulose (999 and 042) and cottonseed hulls (937 and 965)


310


Table XIV. Observations of Sheep on Digestion-Metabolism Trials to Determine the Value of H C and Molasses in a Maintenance Ration

Av. daily gain, gm. Feed effic, gm./gm. gain" Time to change to ration, days Digestible energy, % of gross Digestible protein as %

Basal

Basal + Molasses

Basal + HC

92.10 17.32 5.30 60.90 64.90

73.40 16.45 4.80 59.20 61.30

86.10 15.16 4.80 58.50 58.50


° Not adjusted for DM content of rations. animals feed on shrubs and trees, or on associations in which shrubs and trees play an important part, than on true grasses or grass, legume pastures, short and tall grass ranges, and steppes" (27, 28,29). Perhaps we are either trying to use the wrong woods or have not properly treated or supplemented it. Limitations of previous rations are probably due in part to an inadequate mineral supply. W i t h our current knowledge of the more detailed mineral requirements and mode of interactions, it is possible to provide the basic supplements which are needed for wood derived rations. Of particular interest are the relatively low levels of potassium, calcium, and some of the trace elements in wood products (39). The second limitation in the use of wood is that microbial fermentation of wood products is slow. W h i l e it has been shown that Solka Floe (trade mark of the Brown Paper Co., Montreal, Quebec) an£ filter paper powder can be completely digested if exposed to rumen fluid for 72 hours, the particle size of these is such that they are likely to pass from the Table X V .

Composition of Rations and Coefficients

DM Ration 07 39 41 01 03 40 42 02 04 29

Basal High Fiber + 15% L H C + 15% N L H C + 15% U H C + 15% N U H C + 30% L H C + 30% N L H C + 30% U H C + 30% N U H C + 50% L H C

%

90.2 90.1 89.8 90.3 89.9 90.3 89.4 91.5 91.2 70.9

1 Basal Low Fiber 90.8 2 + 15% Dried L H C 91.6 3 + 30% Dried L H C 91.9

E.E.

COD

%

a

± ± ± ± ± ± ± ± ± ±

Fiber

COD'

± 1.3

3.2 1.1 3.4 2.0 2.3 1.0 0.7 1.3 1.0 1.4

0.9 0.8 0.7 0.6 0.6 0.7 0.5 0.7 0.6 1.1

76.0 73.4 66.8 60.9 48.9 57.0 45.4 71.3 62.0 66.4

0.4 92.5 87.5 ± 1.9 81.3 ± 2.1

3.8 4.8 3.5

90.9 0.8 88.5 ± 1.6 84.3 2.4

40.1 47.6 46.1 42.0 45.5 52.6 54.7 50.6 49.5 51.8

± ± ± ± ± ± ±

COD'

%

0.5 4.8 1.4 9.2 3.2 0.8 6.6 4.6 6.6

± ± ± ± ± ± ± ± ± ± ± 1.8 69.2 1.9 38.0 ± 2.2 35.4 ±

38.7 33.5 31.6 31.8 30.9 27.7 26.7 26.2 27.2 14.5

46.5 42.8 37.1 34.2 36.3 40.1 40.5 34.3 36.2 21.3

• Mean ± standard deviation.


3.1 1.2 3.9 2.3 2.2 2.0 1.2 1.2 2.1 0.8 4.7 13.2 14.7


18.

PFANDER E T A L .

311

Ruminant Nutrition

rumen before the complex hydrolytic processes are completed. Thus, most of these fine particles escape before being digested. The opportunity seems to present itself for developing a tailor-made roughage product from wood residues. Such products could either be inert or have relatively high nutritional value, depending upon the economics of production, distribution, and the comparative prices of competitive products. The use of the hemicellulose fraction of wood offers great promise. Some of the results reported herein have been reported in preliminary reports (43) and confirmed in part in other species (1, 4, 8, 39, 42). Hemicellulose can be ammoniated (37). Other wood by-products have been tested with generally favorable results (18). Other reports of this symposium cover these aspects. One should be alert to the possible toxicity of products fed at high levels. Such toxicity could be caused by other extractives from wood or to substrate inhibition. Repolymerization may occur if the products are heated (Table X V ) . However, our trials indicate no toxicity at generally acceptable levels of carbohydrate addition, and the dried product may serve as a roughage replacement. A basic problem in ruminant nutrition which still needs additional work is the area of adaptation to rations and the possibility of providing specific microorganisms to handle the components in the mixture. W h i l e the addition of starch to ruminant rations has been shown to cause a marked shift in bacterial populations (26) similar information is missing on the build-up and adaptation to the addition of other carbohydrates. Highly soluble carbohydrates generally lead to a much more acid rumen and to the destruction of a number of protozoa. Fermentation patterns of Digestibility of Rations Used i n Trials 2 & 3 Crude Protein %

9.2 9.1 9.5 8.4 8.8 9.2 8.9 8.8 8.6 6.3 14.3 12.8 14.7

COD*

Ash

NFE

%

%

1.9 1.2 3.3 0.8 1.3 0.9 2.4 1.1 5.8 3.0

4.4 4.0 4.7 3.8 4.2 4.0 5.4 3.9 4.8 5.3

37.0 42.7 43.3 45.7 45.4 48.7 47.9 51.9 50.0 43.7

87.4 ± 0.7 78.7 ± 4.1 75.2

3.2 2.6 3.6

67.8 69.6 67.9

42.0 42.7 48.4 42.5 43.7 52.5 53.0 47.2 42.5 41.9

± ± ± ± ± ± ± ± ± ±

TDN

GE

%

COD

COD

a

31.2 52.4 51.5 47.6 52.1 60.0 62.3 59.6 57.0 62.1

± 4.6 ± 1.9 ± ± ±

3.1 2.2 3.3 1.2 1.0 1.7 1.0 1.8

± 95.5 ± 0.3 91.7 ± 1.3 85.4 ± 1.7

34.9 41.9 39.7 37.1 39.3 46.1 45.9 45.2 43.0 34.6

± ± ± ± ± ± ± ± ± ±

a

2.8 1.1 2.8 1.8 2.0 0.9 0.6 1.3 0.9 l.l

47.0 45.6 40.7 41.8 51.5 53.2 49.4 48.8 f t

86.9 ± 0.3 84.7 ± 1.7 76.5 ± 1.8

* Equivalent to 44% TDN on 90% dry matter basis.


± ± ± ± ± ± ± ±

1.0 2.9 2.6 1.8 0.8 1.1 1.1 0.9

312


Table XVI.

Analysis of Variance of the T D N in Hemicellulose Rations

Source

DF

SS

MS

F

Sheep Area (Ar) Acid (AC) Level (L) Ar x L Error Total

3 1 1 1 1 23 30 •

2.13 41.63 3.19 243.65 30.01 59.18 379.79

41.63 3.19 243.65 30.01 2.57

16.20

6

94.80 11.68* b

• One degree of freedom used to calculate missing value. p < 0.01. Downloaded by UNIV OF GUELPH LIBRARY on June 8, 2012 | http://pubs.acs.org Publication Date: June 1, 1969 | doi: 10.1021/ba-1969-0095.ch018

6

may also be changed. Thus, it appears that the organisms involved are constantly available in the population and it merely takes the presence of large amounts of starch to induce the changes. Cellulolytic organisms possibly require more negative oxidation-reduction potentials than those produced by the average ration. Some artificial means of reducing redox potentials might result in a better adaptation of rumen microorganisms. Summary The literature reviewed indicates that raw wood, as browse, and products such as leaves and beans can be used in ruminant rations, and may be the most important energy source for animals. Sawdust and other raw woods are poorly utilized as energy sources but may be used at the 10% level to replace hulls or roughage i n high grain rations. Cellulose prepared from wood is digestible, but particle size limits utility. W o o d molasses and other hydrolytic products are well digested. Derived products containing hemicellulose were shown to be palatable and to provide utilizable energy for ruminants. Hemicellulose from various sources and treated by various methods was calculated to yield from 1800 to 3100 kcal. D E / k g . The drying of these products apparently formed some polymer which reduced the utility of the ration at high levels. The possibility of toxic effects when rations are supplied at the 15% level appears remote. Hemicellulose when supplying over 30% of ration dry matter and when dried or neutralized reduces palatability and lowers digestibility. Acknowledgments This paper is a contribution from the Missouri Agricultural Experiment Station, Columbia, Missouri and U . S . D . A . Forest Products Laboratory, Madison, Wisconsin.


18.

PFANDER E T A L .

Ruminant Nutrition

313


The authors acknowledge the technical assistance of: Orval Lewis in care of management of experimental animals; Wayne Reichert i n bomb calorimetry; C . W . Gehrke and staff for A O A C analysis; Harold Hedrick and staff for carcass data; R. L . Preston for computer programs used i n data analysis; J. F . Saeman and staff for sugar analysis; Dale Galloway, Dale Turner, and T . R. Edgerton for many fruitful discussions leading to the designs of some of the trials reported. The projects reported were supported i n part by grants from Masonite Corporation, Chicago, Illinois. Supplies used were furnished b y International Minerals, Chicago, Illinois; Calcium Carbonate Corpora tion, Quincy, Illinois; Hoffman L a Roche, Nutley, N e w Jersey; Dawes Laboratory, Chicago, Illinois; and Thompson-Hayward, Kansas City, Missouri. Literature Cited (1) Algeo, J. W., Brannum, T. P., Hibbits, A. G., Proc. Western Sec. Am. Soc. AnimalSci.,19th, Santa Ynez, California (1968). (2) AOAC, "Official Methods of Analysis," Washington, D.C.,1965. (3) Archibald, J. G., Massachusetts Exp. Stat. Bull.230,160(1926). (4) Bartley, Ε. E., Farmer, E. L., Pfost, H. B., Dayton, A. D., J. Dairy Sci. 51(5), 706 (1968). (5) Bergius, F.,J.Soc. Chem. Ind.52,1045(1933). (6) Bonsma, J.C.,Farming in S. Africa 17, 226 (1942). (7) Ibid.,17,259(1942). (8) Boren, F. W., Pfost, H. B., Smith, E . F., Richardson, D., Kansas Agr. Exp. Sta. Bull. Β 483 (1965). (9) Breirem, K., Papirjournalen28,101(1940). (10) Ibid.,28,113(1940). (11) Ibid.,28,119(1940). (12) Ibid., 28,133(1940). (13) Ibid., 28,152 (1940). (14) Bunger, Landwirtsch. Versuchs-Stat. 126, 1 (1936). (15) Burkitt, Wm. H., Lewis, James K., Van Horn, J. L., Willson, Fred S., Agr. Exp. Sta. Bull. 498, Bozeman, Montana (1954). (16) Cody, R. E., Jr., Morrill, J., Hibbs, C. M., J.DairySci.51 (6), 952 (1968). (17) Colovas, N. F., Keener, Η. Α., Prescott, J. R., Teeri, A. E., J. Dairy Sci. 32, 907 (1949). (18) Cullison, A. E., Ward, C. S., J. Animal Sci. 24, 877 (1965). (19) De Souza, A. J., Bol. Minist. Agr. Brasil 32, 1 (1943). (20) Ellis, W. C., M.S. Thesis, Univ. of Missouri, Columbia, Missouri (1955). (21) Hargus, W. Α., Ph.D. Thesis, Univ. of Missouri, Columbia, Missouri (1962). (22) Herbst, Walter, U. S. Patent 2269665 (1936). (23) Holtan, E. M., Holtan, M. D., Papirjournalen 28, 157 (1940). (24) Honcamp, F., Hilgert, H., Wohlbier, W., Biochem. Ztschr. 248, 474 (1932). (25) Hvidsten, H., Homb, T., Proc. 11th Intern. Congr. Pure Appl. Chem., London, England 3, 13 (1947). (26) Hungate, "The Rumen and Its Microbes," Academic Press, New York, 1966.



314

CELLULASES A N D THEIR

APPLICATIONS

(27) Imperial Agricultural Bureaux, Imp. Agr. Bur. Joint Publ. No. 10, viii (1947). (28) Ibid., No.10,231(1947). (29) Ibid., No. 10,xxxiv(1947). (30) Informal Committee on Forage Evaluation, unpublished data (1965). (31) Jarl, F., Kgl. Lantbruksakad. Tidskr. 82, 57 (1943). (32) Jones, I.R.,Oregon Cir. 181 (1949). (33) Kellner, O., "The Scientific Feeding of Animals," The Macmillan Company, New York, N. Y., 1913. (34) Kesler, E. M., Chandler, P. T., Branding, A. E., J. Dairy Sci. 239 (1967). (35) Keyes, E. A., Montana Cir. 202 (1953). (36) Lyons, D. T., M.S. Thesis, Univ. of Missouri, Columbia, Missouri (1958). (37) Magruder, N. D., Knodt, C. B., J. Dairy Sci. 36, 581 (1953). (38) Mangold, E., Bruggemann, H., Theel, E., Landwirtsch. Jahrb. 78, 649 (1933). (39) Moore, J. D., M.S. Thesis, Univ. of Oklahoma, Norman, Oklahoma (1968). (40) Morrison, F. B., "Feeds and Feeding," 22nd Ed., The Morrison Publishing Company, Ithaca, N. Y., 1956. (41) Peet, H. S., Ragsdale, A. C., Missouri Agr. Exp. Sta. Bull. 605 (1953). (42) Perry, T. W., Hiller, R. J., Shepard, J. P., Beeson, W. M., Purdue Agr. Exp. Sta. No. 14, Re. Prog. Rep. 153 (1964). (43) Pfander, W. H., Ross, C. V., Preston, R. L., Vipperman, P. E., Tyree, W., Univ. of Missouri, Suppl. Special Rep. Bull. 38 (1964). (44) Poijarvi, I., Acta agral. fenn. 57, No. 1, 1 (1944). (45) Preston,R. L., Kruse, C. G., unpublished data (1968). (46) Saeman, J. F., Moore, W. E., Mitchell, R. L., Millett, M. A., Tappi 37 (8) (1954). (47) Stahl, Ztschr. Schweinezucht 43,471(1936). (48) Stephenson, J. N., Ed., "Pulp and Paper Manufacture," Vol. 1, 3rd ed., McGraw-Hill Book Company, Inc., New York, N. Y., 1950. (49) Virtanen, A. I., Koistinen, O. A., Svenskkem.Tidskr. 56, 391 (1944). (50) Wisconsin Agr. Exp. Sta. Bull. 323, Annual Report (1920). (51) Woodward, T. E., Converse, H. T., Hale, W. R., McNulty, J. B., USDA Bulletin 1272,9,1924. RECEIVED October 14, 1968. Journal Series No. 5479. Approved by Director (College of Agriculture) Univ. of Missouri.


Cellulases and Their Applications

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