A STUDY OF THE LIGNIN, PROTEIN, AND FAT CONTENTS OF SEVERAL COMMON PASTURE MIXTURES AND THEIR EFFECT ON BEEF PRODUCTION

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A STUDY OF THE LIGNIN, PROTEIN, AND FAT CONTENTS OF SEVERAL COMMON PASTURE MIXTURES AND THEIR EFFECT ON BEEF PRODUCTION

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A STUDY OF THE LIONIN, PROTEIN, AND FAT CONTENTS SEVERAL COMMON PASTURE MIXTURES AND THEIR EFFECT ON BEfcE PRODUCTION

A Thesis

Submitted to the Faculty

of

Purdue University

by

Louis Neal Wise

In Partial Fulfillment of the Requirements for the Degree

of

Doctor of Philosophy

June, 1950

P U R D U E U N IV E R SIT Y

T H IS I S TO CERTIFY THAT TH E T H E S IS PREPA RED UNDER MY SUPERVISION

BY

E N ~X'd.'.L-fcm

rl

■— >

I.■.

a_.a. ._ I. .

t

*J ^

. I.

C O M P L IE S WITH T H E UNIVERSITY REGULATIONS ON GRADUATION TH ESES

AND I S APPRO V ED BY ME AS FULFILLING THIS PART O F T H E REQUIREMENTS

F O R T H E DEGREE OF

^ f (

^ .A* L-'

.

/

.

l/

PH o r K s s o n in

IIf-a d o r S c

C h a r o e o f T h e h is

hool

or

D

epahtmbnt

TO T H E LIBRA RIA N:----IS T H I S T H ESIS IS NOT TO B E REGARD ED AS CONFIDENTIAL.

C.

rBoraaioB nr

GRAD.

SCHOOL

FORM

• —3 . « C —1M

o h a i o b

ACKNOWLEDGMENTS

The author wishes to express his appreciation and to acknowledge his indebtedness to Dr. G. 0. Mott for his sincere interest, assistance, during his entire graduate work.

and encouragement

He is likewise in­

debted to Dr. W. M. Beeson, Dr. G. A.. Gries, Dr. A. Olhrogge,

and Professor 3. R. Miles.

The author extends grateful thanks to T. A. Dykes, J. Martin,

and others who assisted in many ways*

TABLE OF CONTENTS Page ....................................................

i

I NTRODU C TI O N ...............................................

1

SURVEY OF L I T E R A T U R E .....................................

5

P R O C E D U R E ..................................................

33

RESULTS AND D I S C U S S I O N

...................................

45

Results of the Chemical Analysis of Ladino Clever, Bromegrass, end Ladino Clover Bromegrass Mixture at Two-Week Intervals ......

47

...................................

1+7

ABSTRACT

L a d i n o C lover

Bromegrass

..................................

Ladino Clever - BromegrassMixture

51

.......

54

Results of the Chemical Analysis of Alfalfa, Timothy, and Alfalfa - Timothy Mixture at Two-Week Intervals ..........................

59

A l f a l f a .......................................

59

......................................

62

Timothy

Alfalfa - T i m o t h y M i x t u r e .................. Results of the Chemical Analysis of Birdsfoot Trefoil, Kentucky Bluegrass, and BIrdsfcot Trefoil - Kentucky Bluegrass Mixture ........ Birdsfoot Trefoil

61+

63

..........................

68

........................

71

Birdsfoot Trefoil - Kentucky Bluegrass Mixture ....................................

73

Results of the Chemical Analysis of Birdsfoot Trefoil, Alta Fescue, and Birdsfoot - Alta Fescue Mixture ........................

77

Kentucky Bluegrass

Birdsfoot Trefoil Alta Fescue

.........................

77

................................

79

Birdsfoot Trefoil - Alta Fescue Mixture

82

TABLE OF CONTENTS - Continued Page RESULTS AND DISCUSSION - Continued Results of the Chemical Analysis of Forage from a Permanent Bluegrass Pasture Receiving 60 Pounds PgOfJ* 30 Pounds K 2O, and 60 Pounds of Nitrogen A n n u a l l y .........................»• Permanent Bluegrass Mixture

...............

Results of the Chemical Analysis of Forage from a Permanent Bluegrass Pasture Receiving 60 Pounds P2O5 and 30 Pounds KgO Annually ••••••• Permanent Bluegrass Mixture

..............

The Relationships Existing Between the Various Chemical Components and Between Certain of these Constituents and the T.D.N. Produced Per Pasture forthe Mixtures Studied •..••••••

87 67

91 91

96

SUMMARY AND C O N C L U S I O N S ...............................

118

B I B L I O G R A P H Y ..........................

122

A P P E N D I X ...............................................

131

LISTS OF TABLLS AMD FIGURES

List of Tables Table 1.

Page Seeding Rates and Annual Fertilization of Rotationally Grazed Pastures at the Miller-Purdue F a r m ........................

35

2.

Comparative C lima to logical Date in Indiana ••

l\2



Species Found at the Miller-Purdue Farm

I4.3

if*

5#

6.

Chemical Composition of Ladino Clover at Two-Week Intervals Throughout the Grazing Season ..................................... *

I4.S

Chemical Composition of Bromegrass at TwoWeek Intervals Throughout the Grazing Season



52

Chemical Composition Throughout the Grazing Season of Forage from Ladino Clover Bromegrass Pasture and the Total Di­ gestible Nutrients Produced Per Pasture • •

55

7.

Chemical Composition of Ladino Clover, Bromegrass, and Ladino Clover - Brome­ grass Mixture----Average of Three TwoWeek Periods ......................... .

8*

Chemical Composition of Alfalfa at Two-Week Intervals Throughout the Grazing Season *•

60

Chemical Composition of Timothy at Two-Week Intervals Throughout the Grazing Season • •

63

Chemical Composition Throughout the Grazing Season of Forage from Alfalfa - Timothy Pasture and the Total Digestible Nutrients Produced Per Pasture ..............

65

Chemical Composition of Alfalfa, Timothy, and Alfalfa - Timothy Mixture----Average of Three Two-Week Periods *•••.•..••.••••*

6?

9« 10*

11.

12*

Chemical Composition of Birdsfoot Trefoil at Two-Week Intervals Througnout the Grazing Season ........................ .

List of Tables - Continued Table 13•

l4»

15.

16.

17.

18*

19*

20.

21.

22. 23.

Page Chemical Composition of Kentucky Bluegrass at Two-Week Intervals Throughout the Grazing Season .....

72

Chemical Composition Throughout the Grazing Season of Forage from Birdsfoot Trefoil Kentucky Bluegrass Pasture and the Total Digestible Nutrients Produced Per Pasture •

74

Chemical Composition of Birdsfoot Trefoil, Kentucky Bluegrass, and Birdsfoot Trefoil Kentucky Bluegrass Mixture----Average of ............... Three Two-Week Periods

76

Chemical Composition of Birdsfoot Trefoil at Two-Week Intervals Throughout the Grazing Season ................................

78

Chemical Composition of Alta Fescue at Two7/eek Intervals Throughout the Grazing Season ....................................

80

Chemical Composition Throughout the Grazing Season of Forage from Birdsfoot Trefoil Alta Fescue Pasture and the Total Digest­ ible Nutrients Produced Per Pasture •••••••

63

Chemical Composition of Birdsfoot Trefoil, Alta Fescue, and Birdsfoot Trefoil - Alta Fescue Mixture— --Average of Three TwoWeek Periods ......................

86

Chemical Composition of Forage from Permanent Bluegrass Pasture Receiving 60 Pounds P2O5 # 30 Pounds KpO, and 60 Pounds of Nitrogen Annually ana the Total Digestible Nutrients Produced Per Pasture .................

88

Chemical Composition of Forage from Permanent Bluegrass Pasture Receiving 60 Pounds P2°5 and 30 Pounds KoO Annually and the Total Digestible Nutrients Produced Per Pasture •

92

Chemical Composition of Permanent BluegrassNPK Average of Three Two-Week Periods ••

95

Chemical Composition of Permanent BluegrassPK----Average of Three Two-Week Periods •••

95

List of Tables - Continued Table 2lj.*

25*

26.

Page The Relationships Existing Between the Various Components Determined for Ladino Clover •••••••••••••••••.................

96

The Relationships Existing Between the Various Components Determined for Bromegrass • • ..... ...........

99

The Relationships Existing Between the Various Components Determined for Ladino Clover - Bromegrass Mixture

100

27 •

The Relationships Existing Between the Various Components Determined for Alfalfa •101

28.

The Relationships Existing Between the Various Components Determined for Timothy •102

29.

The Relationships Existing Between the Various Components Determined for Alfalfa Timothy Mixture .... .

103

The Relationships Existing Between the Various Components Determined for ..................... Birdsfoot Trefoil

IOI4.

The Relationships Existing Between the Various Components Determined for Kentucky Bluegrass *••»............

105

The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil - Kentucky Bluegrass Mixture ....••

106

The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil *••••••.••....*•••••...............

107

The Relationships Existing Between the Various Components Determined for Alta Fescue ...........................

10 8

The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil - Alta Fescue Mixture ........

109

The Relationships Existing Between the Various Components Determined for Permanent Bluegrass Mixture - NPK . ............... *

110

30*

31*

32.

33.

34*

35*

36.

List of Tables - Continued Table 37.

38.

Page The Relationships Existing Between the Various Components Determined for Permanent Bluegrass Mixture - PK ...................

111

Compilation of All Correlation Coefficients Found i n Tables 1-1if .............

113

Appendix Tables 1. 2. 3.

if. 5* 6. 7. 8. 9.

10. 11.

Detailed Data on the Chemical Ladino Clover - Percent

Composition of ........••••

131

Detailed Data on the Chemical Composition of Bromegrass - Percent .....................

132

Detailed Data on the Chemical Ladino Clover - Bromegrass cent

133

Composition of Mixture - Per­

Detailed Data on the Chemical Composition of Alfalfa - Percent ...........................

134-

Detailed Data on the Chemical Composition of Timothy ......................................

135

Detailed Data on the Chemical Composition of Alfalfa - Timothy Mixture - Percent .......

13&

Detailed Data on the Chemical Composition of Birdsfoot Trefoil • ••••.................

137

Detailed Data on the Chemical Composition of Kentucky Bluegrass - P e r c e n t ...............

13$

Detailed Data on the Chemical Composition of Birdsfoot Trefoil - Kentucky Bluegrass Mixture - Percent .......................

139

Detailed Data on the Chemical Composition of Birdsfoot Trefoil - Percent ............

lifO

Detailed Data on the Chemical Composition of Alta Fescue - P e r c e n t .......................

1^1

List of Tables - Continued Ap pe n d i x

Table 12*

13« 14.

Page Detailed Data on the Chemical Composition of Birdsfoot Trefoil - Alta Fescue Mixture .......... Percent

llp2

Detailed Data on the Chemical Composition of Permanent Bluegrass N-P-K ........

143

Detailed Data on the Chemical Composition of Permanent Bluegrass P-K .......

145

List of Figures Figure 1*

Page Arrangement of Pasture Plots Miller-Purdue Farm .......................

30

ABSTRACT

There were two principal phases of this investiga­ tion.

One was a study of the seasonal changes in the

chemical composition of several pasture mixtures and of the principal grasses and legumes making up each of the mixtures.

The second phase dealt with the establishment

of relationships between the chemical constituents of each mixture and species, and between these constituents and the production of total digestible nutrients. The study was made on pastures at the Miller-Purdue Farm and in conjunction with grazing trials which have been conducted there for several years.

Six pastures or treat­

ments were considered; there were four grass-legume mix­ tures and two permanent bluegrass pastures. received 60 pounds ^2^5

30 pounds of KgO annually.

One of the permanent pastures received, pounds of nitrogen. three times.

All pastures

in addition, 60

Each of the six pastures was replicated

All were rotationally grazed.

Each field was divided into three paddocks.

The

cattle, Hereford steers, remained on each paddock for lip days, after trialch they were weighed and transferred to the next paddock.

There were twelve llp-day periods.

Three forage sample were obtained prior to the be­ ginning of each grazing period.

One was a mixture sample

which contained principally the sown species but might

contain any other species of plant which infested the pas­ ture*

The other two samples consisted of the sown grass

and legume, carefully separated to give a pure species of each* A chemical analysis was made of the forage samples to determine:

dry matter, crude protein, lignin, ether ex­

tract, and a combination of digestible protein and soluble carbohydrates designated as "Fraction X"* Dry matter was found to be unusually high in the spring but did increase to a seasonal peak in early or mid­ summer* fall*

It then dropped to its lowest point in the late The dry matter content of legumes was much lower

than that of grasses, with the forage mixtures generally intermediate* The lignin content showed a trend similar to the dry matter*

The legumes again were lowest.

Kentucky bluegrass

showed the highest lignin content of any species or mixture but was closely followed by the forage mixture from the permanent bluegrass pasture which did not receive the nitrogen* Crude protein was highest in the spring and fall, declining to a seasonal low during the summer months*

At

no period during the grazing season did any of the grasses, legumes, or mixtures show a crude protein content which was less than the minimum requirement for maximum produc­ tion by beef animals*

Ether extract failed to show the definite seasonal trends which were characteristic of the other constituents* This was pro bably due to the small

amount of this fraction

present in forage plants and to the heterogeneous nature of the various substances making u p the ether extract.

It

was highest in the spring and fall periods. As would be expected,

the trend shown by fraction

X closely parall e le d that previously described for crude protein.

The fraction X content was considerably larger

than that of crude protein since this tained the

substance also con­

soluble carbohydrates*

Correlation

end regress i on coefficients and co effi­

cients of determination were used to express the relation­ ships between the seasonal

variation in the chemical con­

stituents and b et w ee n the chemical constituents and the total digestible nutrients produced per pasture* Lignin appeared to be positively correlated with dry matter raid negatively correlated with crude protein, extract and fraction X* large amounts of legumes,

Tnree pastures,

etner

which contained

showed a positive relationship

between lignin end total digestible nutrients. not a causal effect bi’t resulted

This was

from the associative

effect of both lignin and total digestible nutrients with tne dry matter content.

Dry matter ratner than lignin

appeared to be tne limiting factor In beef production on these [pastures.

The three pastures consisting principally

of grass,

showed an inverse relationship between lignin and

total digestible nutrients.

Apparently the dry matter con­

tents of these three pastures were not limiting and lignin was an important factor affecting beef production* The dry matter content appeared to be negatively re­ lated to crude protein, ether extract, and fraction X* The pastures previously referred to which showed a positive correlation between lignin and total digestible nutrients, also showed a positive correlation between dry matter and total digestible nutrients.

For the other pastures there

was an Inverse relationship* Crude protein was positively related to ether extract and fraction X but negatively correlated with total digest­ ible nutrients*

This latter relationship could be expected

since the protein content was never a limiting factor*

A STUDY OF THE LIGNIN, PROTEIN, AND FAT CON T E N T S OF SEVERAL COMMON PASTURE MIXTURES AND THEIR E F F E C T ON BEEF PRODUCTION

INTRODUCTION

Classes of livestock differ as to the extent of their use of forage.

The quantity used by poultry is small but

its importance to them as a source of vitamins, minerals, and protein is undeniable.

Swine are larger consumers of

pasturage but again it is the nutritive balance,

rather

than total consumption, which is of prime importance. Ruminants may, and often do, receive all of their feed in the form of forages and it is this group of livestock which receives, at least quantitatively, the greatest benefit from pastures. Because of the diversity of aims when considering utilization of pasture by the different classes o f live­ stock, further discussion will be limited to ruminants. To insure a successful pasture-livestock enterprise, several conditions must be met#

There must be a n adequate

supply of forage, palatability should be such as to insure sufficient consumption, and the quality and digestibility should be of a high order#

Failure of a pa rt ic ul ar pasture

species or mixture to meet any one of these requirements would seriously reduce i ts value#

2

The vast majority of experimental work has been con­ cerned with increased production of forage.

There is gen­

eral agreement on the more important factors affecting quantity; adequate fertilization, moisture, and proper tem­ perature are standard prerequisites for increasing yields* Data on palatability are much more varied and con­ troversial.

Lists of pasture species in order of prefer­

ence by certain classes of livestock often appear to com­ pletely contradict one another.

The importance of palat­

ability has been questioned, and with considerable justifi­ cation. Consideration of the quality and digestibility of forage must necessarily be made concurrently.

Quality in­

volves the balance of the various plant constituents present digestibility denotes their availability to the animal*

For

the most exacting information on digestibility, digestion trials are necessary.

Due to the difficulties Involved in

conducting such trials with grazing animals, the availabil­ ity of a nutrient is normally assessed on the basis of standard digestion coefficients.

These coefficients do not

take into consideration the rather significant seasonal changes which take place as a result of increased plant maturity* Experimental evidence shows a definite negative corre­ lation between the lignin content of feed and subsequent animal gains.

It has been suggested that lignification may

be the most important factor limiting animal productivity* It should be noted, however, that this reverse relationship between lignin and gain has been obtained from experiments Involving a known or controlled feed intake.

Although

there is little doubt that a similar relationship exists In practical grazing trials,

It is much more difficult to

establish proof. Experimental data presented by Forbes and Garrigus indicate a straight line negative relationship between lignin content and dry matter digestibility.

Yet they

stated that such a relationship cannot be assumed for for­ ages other than those tested and that data must be accumu­ lated for a large number of forages under a variety of con­ ditions and in trials with the species of animals to which the data are to be applied.

These investigators suggested

that with such Information available practical regression equations may be established for predicting nutritive value of growing pasture forage from the lignin content alone. All classes of livestock require some fat in their diet.

This requirement arises not only from the essenti­

ality of certain fatty acids, but from the numerous roles played by other non-fat constituents of the lipid group. These include the all important phospholipids, fat soluble vitamins.

sterols, and

The importance of fat as a source of

total digestible nutrients cannot be over looked since It contains 2.25 times the energy content per unit weight as do

carbohydrates and proteins.

The significance of this high

energy value may be modified by the somewhat lower digest­ ibility of a large portion of the lipids. The role of proteins in the nutrition of animals is well known, and much emphasis has been placed on the value of increasing the protein content of our forages.

The

importance of supplying to the grazing animal its minimum protein requirement is undeniable.

Caution should be exer­

cised, however, to avoid misplaced emphasis on the impor­ tance of that portion of the protein content which is above the minimum requirement of the animal.

The quality of pro­

tein which is so vitally important for non-ruminants, may be of little consequence in the nutrition of ruminants. The present study will attempt to trace the seasonal variation in the protein, fat, and lignin content of several of the more common pasture species and mixtures.

The Inter­

relationship existing between these constituents will be studied and the role of each evaluated on the basis of the performance of grazing animals. be considered.

Total production will not

SURVEY OF LITERATURE

Initial work on lignin appears to have been brought about by widespread dissatisfaction with nitrogen free ex­ tract and crude fiber as measures of the carbohydrate frac­ tion of plants.

Nitrogen free extract has been assumed to

be composed of sugar, starches, and other easily soluble carbohydrates, ible.

and as such, to be almost completely digest­

Crude fiber, on the other hand, has been classed as

the more or less indigestible fraction.

Nutritionists and

biochemists agree that the digestibility of carbohydrates does not consistently follow this separation. The Wiende method for the determination of crude fiber has been in general use with only minor alterations since l88ip (26).

It involves the hydrolysis of the plant material

with dilute acid and alkali.

Heller and Wall (l\S) point out

that it fails to give a true conception of the chemical com­ position of the residue.

Consequently, there is no way of

estimating the relative digestibility of the quantity of crude fiber reported.

Sullivan and Garber (88) report that

preparations of crude fiber from various types of forages contained on the average 85 percent cellulose and 12 percent lignin.

However, not all of the lignin and cellulose were

present in this fraction.

Crude fiber was found to contain

only Ip—6»7 percent of the total plant lignin and l±0-68 percent of the cellulose.

The chemical make-up of crude fiber from

various forages, or a single forage at different stages of maturity, may be quite different. Nitrogen free extract is not determined directly but is obtained by the difference method.

The errors inherent

in the crude fiber determination will be reflected in this fraction.

By definition the nitrogen free extract repre­

sents "easily soluble carbohydrates•M

Actually it contains

certain indigestible substances which were dissolved out of the crude fiber (ip6>) (B 6>) •

When only ip percent of the plant

lignin is contained in the crude fiber, 9 & percent is to be found in the nitrogen free extract.

Stanley and Hodgson

(8 6 ) concluded tnat tne onset of lignification during later stages of growth may explain in part the increase in nitro­ gen free extract which is often concurrently observed. It is apparent that the division of carbohydrates into crude fiber as the poorly digestible fraction, and nitrogen free extract as the highly digestible fraction, justifiably questioned.

can be

Crampton and Maynard (20) have sug­

gested a modified procedure for feed analysis which makes a sharper distinction between carbohydrates with respect to their digestibility.

They propose to diviae tne carbony-

drates into three fractions: portion,

a practically indigestible

li0nin; a highly digestible fraction, otner carbo-

nydrates; ana cellulose,

tne digestibility of whicn may be

expected to vary inversely witn lignification.

Other in-

vesti0ators (2 2 )(68)(6 9 ) have found tnis modified procedure

to be superior to the conventional method of feed analysis. The exact chemical structure of lignin is not known. Phillips (77) reports that the presence of the methoxyl group (-OCH^) has been definitely established. groups are difficult to split out,

Since these

they are probably

attached in the form of an ether linkage.

The number of

groups varies with the source and method of isolation of the lignin.

There is abundant evidence (24) (77) to support

the claim that lignin, unlike cellulose, structure.

is cyclic in

The presence of hydroxyl groups in lignin is

indicated by the fact that it can be acetylated or alkylated. Lignin is generally considered not to be a single com­ pound but a combination of several substances of a similar organic nature (79)(27)(62)(6 3 )•

Freudenberg (34) found

that lignin exists in wood in different degrees of condensa­ tion, ranging from simple units of low molecular weights to highly complex aggregates.

There is no reason to believe

that a similar situation does not exist in forage plants. Lignin occurs in plants chiefly as a lignocellulose (20).

Tne exact mechanism of the deposition of lignin is

unknown.

The basic unit making up the cell wall of plants

is the long chain-like cellulose molecule.

Tnese cellulose

molecules are aggregated into bundles, or micelles.

Meyer

and Anderson (62) regard the cellulose wall as a structure composed of molecular aggregates or micelles welded together by interlocking cellulose molecules.

Intermicellar spaces

form an i n t e r - c o n n e c t i n g s ys t em b e t w e e n m i c e l l e s . immature p l a n t

these

pectic c o m p o u n d s . placed to v a r y i n g Miller

s paces

are p r i n c i p a l l y f i ll e d with

As the p l a n t matures,

(6 3 ) cites e v id e n c e w n i c h is i n g e n e r a l a g r e e ­

wnich l i g n i f i c a t i o n of l ig n o c e l l u l o s e

si iply a c ha n ge

of

and o t h e r n o n - c e l l u l o s i c

In h e a v i l y

p a t t e r n of is

an

(ip) found

con stituents were de­

and r e s u l t e d in two continuous,

systems.

tnere

Hailey

Inter-communicating, interstices

system m a y be d i s s o l v e d w i t h o u t

dicates

Lignification

tne cellu l os e u ni t s but

in i n t e r m i c e l l a r ma t er i a l .

of the cellulose

structural

He re f er s to the f or m a t i o n

as an i n f i l t r a t i o n pro cess.

posited in t h e elongat e d,

penetrating

e x p l a n a t i o n of the m a n n e r in

takes place.

may not in volve a m o d i f i c a t i o n

lignin

this p e c t i n is r e ­

d e grees w i t h lignin.

ment with M e y e r and A n d e r s o n ' 3

tnat

In an

the other.

inter­

lignified tissue,

eit her

seriously modifying, the .York by

actual chemical

Fhillips

combination

(75)

In­

between

Dijnin and cellulo s e. Tne p r e c u r s o r s o f l i g n i n are l ar gely a m a t t e r of speculations.

Lignified

tissue

con tains relatively large

quantities of l i g n i n and h e r d c e l l u l o s e w i th only a trace of pectin.

T is s u e s w n i c h heve not

u n de r g o n e l i g n i f i c a t i o n co n­

tain r e l a t i v e l y la r ge a m ou n t s of p e c t i n and only treces of h e m i c e l lu l os e ana lignin. by many Norman

as the s u b s t a n c e

Pe ctin has thus b e e n conside r ed from wn i ch lign in

is formed.

(6 7 ) sums up the t he o r i e s involving tr a ns f ormations

of pectin, hemicellulose,

and lignin as follows:

1.

Pectic material is transformed

into lignin.

2*

Hemicellulose may be converted

to lignin.

3*

Polyuronide hemicelluloses are

formed from pectin.

4*

Relationships exist between all three, lignin being formed from pectin through the intermediate stage of polyuronide hemicellulose.

Norman does not cite these relationships as conclusive proof of the manner of formation of lignin, but as convey­ ing an impression of probability.

Sullivan and Garber (8 8 )

point out that the evidence for the above hypothesis was not derived from pasture plants. Bennett

(7)(8 ) working with Kentucky bluegrass and

red clover found definite amounts of cellulose and lignin In the very earliest samples obtained.

This suggests that

these compounds may be active metabolic products rather than products characteristic of maturity in plants.

Approxi­

mately the same percentage of lignin was found to be as­ sociated with different proportions of pectin and hemi­ cellulose in the two species.

Bennett found no direct

evidence for a relationship between pectin, hemicellulose, and lignin. Phillips (76) and associates obtained results which indicated that the plant does not synthesize lignin from cellulose, pectin or pentosans.

They suggest that lignin

Is f o m e d directly from either glucose or sucrose. According

10

to this hypothesis the first step is the production of a substance or substances having firmly bound methoxyl groups* Evidence for this has been the identification of these sub­ stances and their gradual decrease with an increase in the lignin content. Although the present investigation is primarily in­ terested in lignin from the standpoint of its effect upon nutritive value of forage, the role which it plays in the soil cannot be overlooked.

The degree of its susceptibility

to decomposition in the soil is controversial.

Phillips

(77) shows that isolated lignin is very resistant to the action of soil organisms and to wood destroying fungi.

On

the other hand, natural lignin under aerobic conditions can be decomposed by soil microorganisms.

The rate is generally

much slower than for cellulose and hemicellulose. As would be expected,

the decomposition of naturally

occurring lignin is greatly affected by temperature.

At

7 degrees C Phillips (77) found the loss of lignin neglig­ ible whereas at 37 degrees C over 30 percent of the lignin was removed in three and one-half months and 50 percent to 60 percent in nine months.

Under anaerobic conditions

lignin is preserved almost quantitatively.

Waksman and Syer

(9 2 ) concur with these findings but add to the factors which slow down the decomposition of lignin, the condition of in­ creased soil acidity.

Tenny and Waksman (90) show that the

difference in the rapidity of decomposition of the various

chemical constituents under anaerobic conditions are very striking. Waksman and Hutchings (93) point out the specific ef­ fect which lignin exerts upon two of the most important forms of soil nitrogen, namely ammonia and protein*

The

ammonia is absorbed by lignin and held until utilized by plants,

soil microorganisms, or oxidized by specific bac­

teria to nitrate*

The manner in which the proteins combine

with lignin is not fully understood*

However, the complexes

which result from this combination are highly resistant to decomposition and form an essential group of humus constit­ uents*

Fuller (37) indicates that one of the most important

effects of lignin is its depressing action on protein de­ composition*

He cites experimental evidence showing that

the presence of purified lignin in a sandy loam soil def­ initely reduced the amount of nitrate nitrogen recovered from dried blood or ammonium sulfate.

The depressing effect

which lignin had on the decomposition of protein was found by Waksman and Syer (91) not to be the result of a toxic action, but to be an interaction between lignin and protein which results in the formation of a complex "humus nucleus." There have been indications by other workers (6 8 ) that iso­ lated lignin has a bacteriostatic action. Gottlieb and Hendricks (ZfO) call attention to the fact that many investigators have found that as much as 60 to 80 percent of the organic matter in some soils could be ac­

12

counted for as lignin and protein,

They also point out that

it is possible to add acid to an alkaline solution of a pro­ tein and lignin and obtain a precipitate which, resembles extracts of soil organic matter in its empirical properties. Bartlett and Noiroan (5) agree that most of the properties of humus can be explained o n the assumption that it is largely composed of residual lignln from parent plant materials, combined or associated with a nitrogenous complex. Miller and associates

(6 3 ) found a highly significant

correlation between percentage increase in base exchange capacity of decomposing organic matter and percentage in­ crease in lignin content.

Since, however,

the increase in

the base exchange capacity was greater than the increase in the lignin content,

it was suggested that a change takes

place in the absorptive capacity of the lignin as decomposi­ tion progresses. Very recent work by Gottlieb and Geller (/f) states that after a survey of a large number of species of wood rotting fungi, several were found which were capable of m a k ­ ing satisfactory growth on a media in which Isolated lignin was the only source of carbon.

The lignin used was isolated

by a process which they believed mini-ay

12

22.3

ll.il

15.3

3.3

26

29.8

15.3

10.3

2.3

9

37.il

12.9

iii.5

3.5

22

ill.3

lil.7

13.0

3.3

7

36.3

13.5

13.'4

3.4

21

29.0

13. J+

18.0

4.3

k

32.2

13.3

17.3

4.3

18

3lp.l

15.2

I 8.9

il.7

1

29.9

15.9

18.5

4.5

13

29.6

13.3

1 8 .i:.

3.9

28

22.2

ll.l

22.8

ii.o

31.2

13.5

l6,6

3.8

June

July

August

oept •

Average

Least Significant Difference at: $c /o

6*2

l •.5

2.9

1%

3.5

2.0

ii.o

.6

Birdsfoot Trefoil - Kentucky Bluegrass Mixture Table 1!(-•

A considerably lower dry matter content

was found for the mixture than was reported for Kentucky bluegrass alone.

This is due to the modifying influence

of tne extremely low dry matter content of the birdsfoot trefoil.

Part of it may have been the result of the 8.1

percent white clover reported by Dykes to be present in this pasture. Except for the rather high value during the first period,

there Is a progressive increase in the dry

natter content until a seasonal peak is reached in July. Tnis is followed by a general aecline to a low of 18.5 per­ cent in the last period.

The seasonal average was 2ii*9 per­

cent • The average lignin content was 12.1 percent and was higher than that of the other mixtures previously consid­ ered.

The predominating influence

of thehigher

lignin

content of Kentucky bluegrass over

the lower percentage

found in birdsfoot trefoil,

easily seen.

can be

Except for

tne lower lignin contents at either the beginning or the end of the season,

there is no consistent trend.

« high of lip•9 percent,

There is

a low of 9«2 percent.

The average crude protein content evidences the effect of the lower protein content of the grass on the mixture as a whole.

Although tnere is a slight peak of lQ.6 percent

in tne early spring,

the higher protein contents are from

mid-season through the late fall, rising at that time to a seasonal high of 23«5 percent.

714-

Table 11^ Chemical Composition Throughout the Grazing Season of Forage from Birdsfoot Trefoil - Kentucky Bluegrass Pasture and tee Total Digestible Nutrients Froduced Per Pasture _________Chemical Composition - Percent Dry Crude Ether Date______ Matter__ Lignin Protein _ E x t r a c t ___ T._D . N. April

28

29.5

10.7

18.6

4*4

1212

May

12

20.9

9.7

15.8

3.6

1705

26

23.1

13.3

12.0

2.7

803

9

27.5

10.8

17.3

if.O

1288

22

30.3

13.9

15.4

3.8

746

7

31.1

34.9

13.8

3.4

1629

21

19.9

11.8

20.6

4*4

751

4

25.1

13.1

18.7

4.8

1044

18

28.0

12.8

19.1

4.7

651

1

23-9

13.1

20.0

5.1

915

15

21.0

1 1 .u

21.5

5.2

757

28

18.5

9.2

23.5

5.0

74

2lf.9

12.1

18.0

4.3

966

June

July

August

Sept.

Average

Least Slgnlfleant Difference a t : 5%

If.8

3.0

3.1

.9

557.9

1*

6.6

l+.l

I+.2

1.2

759.U

The average ether extract was 4»3 percent.

The per­

iods showing the highest fat content correspond generally to the high periods reported for crude protein. Table 15 gives the chemical composition of both species and of the mixture taken from this pasture.

It condenses

the analysis into 6-week averages and the general relation­ ship already discussed can be more easily discerned.

The

effects of the composition of the grass and legume on that of the mixture can be clearly seen. however,

Caution must be used,

since other species may also exert a considerable

influence on the chemical composition of tne mixture.

76

Table 15 Chemical Composition of Birdsfoot Trefoil, Kentucky Bluegrass, and Birdsfoot Trefoil - Kentucky Bluegrass Mixture Average of Three Two-Week Periods

Periods

DryMat ter

Chemical Composition - Percent Crude Ether Fraction Lignin Protein Extract X BIRDSFOOT TRLFOIL

1st 2nd 3rd 4 th

6 6 6 6

weeks weeks weeks weeks

Average

17.0 13.?

6.4 9.7 9.6 7.8

28.5 2{4-.0 2 4 .? 2 6.0

3.7 3.7 3.5 £.2

53.0 53.2 58.5

1 6 .6

8.4

2 5 .7

3.8

56.3

—— — _ ------- -----

17.9 1 8 .0

6 0 .6

KENTUCKY BLUEGRASS 6 6 6 6

weeks weeks weeks weeks

3.2

1 3 .6 1 8 .1

4.4

19.9

3 .1

3 1 .2

13.5

1 6 .6

27.7 38.3 31.5

.

CO

Average

l/f.9

2 6 .9

12 .2 14.4 lip.1 13.3

CO

1st 2nd 3rd 4-th

-----

BIRDSFOOT TREFC IL - KENTUCKY BLUEGR a SS MIXTURE 1st 2nd 3rd 4-th

6 6 6 6

weeks weeks weeks weeks

Average

2ii.5 26.3 2k*3 21.1 24.9

3*7

12.6 11.2

15.5 15.5 19.5 21.7

4 .8

— — ------- -

5.3



12.1

1 8 .0

4 .3

-----

11.2 1 3 .2

3 .6

—— —

77

Results of the Chemical Analysis of Birdsfoot Trefoil, Alta Fescue, and Birdsfoot Alta Fescue Mixture

The botanical composition of this pasture was as follows: Birdsfoot trefoil Alta fescue

8*4 percent 51•7 percent

Others ij.0.0 percent (white clover, weeds, Kentucky bluegrass, wiregrass, and timothy) The birdsfoot trefoil sown Aith the Alta fescue was 50 percent narrow-leaf and 50 percent broad-leaf*

This is

in contrast to the 100 percent broad-leaf birdsfoot trefoil seeded in the previously discussed birdsfoot trefoil Kentucky bluegrass pasture.

Birdsfoot Trefoil Table l6*

The average dry matter content of birdsfoot

trefoil from this pasture is practically identical to that obtained for tnis same legume sown with Kentucky bluegrass* Note the close similarity of the trends shown in both cases and the extremely low dry matter value shown for each at the seme date in the late fall* The average lignin content, 9*^ percent, is slightly higher than that found for the 100 percent broed-leaf

78

Table 16 Chemical Composition of Birdsfoot Trefoil at Two-Week Intervals Throughout the Grazing Season

Dry Matter

Date

Chemical Composition - Percent Crude Ether Lignin Protein Extract

April

28

20.5

7.2

31.9

4*4

May

12

19.0

6.9

27.5

4-2

26

16.0

8.1

22.14

4.0

9

18.9

8.7

23-3

4.3

22

18.7

10.5

21.6

3.7

14.4

11.5

20.3

3.3

12

16.2

11.1

21.7

3*5

4

17.0

10.5

22.8

4.1

18

15.1

9.1

23.6

3.3

1

17.4

9.8

23.1

3.9

15

llf.l

7*9

2ip»l

4 .0

28

8.9

6.9

25.4

4.0

16.4

9.0

24.0

3.9

June

July

August

Sept.

7

Average

Least Significant Difference at: 5%

3.?

1.6

2.0

•4

lfb

4.3

2.2

2.7

.6

79 birdsfoot trefoil.

This mixture of narrow-leaf and broad-

leaf birdsfoot trefoil does not exhibit the extreme seasonal low shown by the other type.

The trend and the periods at

which seasonal lows and highs occur, however, show a re­ markable similarity. The average crude protein content of the mixed birds­ foot trefoil was 214-.0 percent as compared to 25*7 percent previously shown for the broad-leaf type. of the dry matter content,

As in the case

the differences were quantitative,

and the same trends were apparent.

Ether extract, as usual,

was highest at either of tne seasonal extremes, during the periods of less favorable growth.

and lower

The average

seasonal fat values for the two types of birdsfoot trefoil were 3.8 percent and 3*9 percent.

Alta Fescue Table 17.

Tne average dry matter content was 23*3 per­

cent; this is much lower than that found for bluegrass but still higher than the average reported for bromegrass.

Al­

though the initial period showed a fairly high dry matter content,

the peak occurred in late June.

As in the case of

all species and mixtures considered so far, the lowest value was obtained in the late fall. The lignin content increases very rapidly, rising from an initial value of 7.2 percent to the seasonal peek of ^9»3 percent in only four periods.

The normal decline

ao

Table 1? Chemical Composition of Alta Fescue at Two-Week Intervals Throughout the Grazing Season

Dry Date_____ Matter

Chemical Composition - Percent Crude Ether Fraction Lignin Protein Extract______ X____

April

20

27.6

7.2

20.1

3.9

if9.5

May

12

25.3

9.8

15.6

2.9

ifO.if

26

2I4..5

14.0

12.3

1.9

31.7

9

28*2

Uf.3

11.3

2.5

31.3

22

29.7

12.9

12.4

2.6

29.2

7

20.6

12.k

II4..0

2.9

30.6

12

20.0

9.2

19.6

4.2

1*1.3

22.k

9.3

18.7

k-kr

36.6

18

21.9

10.1

20.2

if.3

14-2.1

1

21.2

8.8

18.8

3.8

if3 .if

15

20.6

7.1

19.5

3.9

50.1

28

17.2

6.1

22.8

If.3

52.14-

23.3

10.1

17.1

3.5

I4. 0 .0

June

July

August

Sept*

if

Average

Least Significant Difference at: 5*

3.9

1.3

2.6

•if

if.3

1%

5.3

1.8

3.5

.6

5.8

during the latter part of the season is disrupted only once, by a slight increase in August.

The average lignin

content of Alta fescue was 10*1 percent, higher than that of bromegrass and timothy but considerably less than the 13*5 percent reported for Kentucky bluegrass.

This higher

value for Kentucky bluegrass is surprising when the physi­ cal nature of these two grasses is examined. has a rough, coarse type of vegetative growth.

Alta fescue In direct

contrast to this is the smooth velvety nature of Kentucky bluegrass.

Evidently the lignin content of a forage cannot

be predicted on the basis of its physical feel or appear­ ance. It should be pointed out that this study involves only the determination of the total lignin content found in the various species and mixtures.

Evidence has been cited in a

previous section that the mode of deposition or the stage of forage maturity may influence tne digestibility of lignin and its effect upon the digestibility of the other constituents.

Total quantity may or may not be the most

important criterion by which to judge the nutritional sig­ nificance of lignin. The average crude protein content was 17*1 percent. This is higher than the 16.6 percent reported for Kentucky bluegrass but much lower than the 21.3 percent shown for bromegrass.

The lower values of both bluegrass and Alta

fescue, as compared to bromegrass, may be partly due to the

comparatively low legume populations with which they are associated.

Ladino clover,

with which bromegrass is as­

sociated, is recognized for its superior associative ef­ fects • Ether extract declined from 3.9 percent in the early spring to an unusual low of 1.9 percent.

From this point

It rises steadily until it reaches 4*4 percent in early August.

It remained

latter part of

at a rather high level during the

the season. Fraction X showed a seasonal

average of i^O.O percent.

Note that this is 20 percent

lower than the corresponding mean given for Ladino clover. This is a large difference when it is considered that this fraction X must necessarily be highly digestible.

The only

other grass for which this substance was determined was bromegrass; its mean value was 43*9 percent.

Birdsfoot Trefoil - Alta Fescue Mixture Table 18.

There appears to be three peaks reached in

the dry matter content.

The first occurs on April 28 and

of course is consistent with that found for practically all other species and mixtures.

It has been previously ex­

plained on the

basis of the lower than normal

rainfall

which occurred

early in the 1949 season.

second and

The

highest peak of 26.5 percent is reached in late June and coincides with the date at which Alta fescue would normally be expected to reach maturity, if it were not grazed or

83

Table 18 Chemical Composition Throughout the Grazing Season of Forage from Birdsfoot Trefoil - Alta Fescue Pasture and the Total Digestible Nutrients Produced Per Pasture Chemical Composition - Percent Dry Crude Ether Matter Lignin Protein Extract

Date

T. d . ;

April

28

2 4 *0

8.9

20.2

4.0

1116

May

12

21*8

8.3

18.5

3.5

1725

26

19.1

11.0

14.2

2.5

856

9

2^.2

10.9

18.0

3.1

1313

22

26.5

10.5

13.7

3.0

959

7

20.6

10.5

14.1

3.2

1258

12

21.0

9.8

18.4

4.4

727

4

21.7

8.4

18.1

4.5

1366

18

23.8

13-6

18.6

4-9

883

1

23.0

14.7

18.0

4.0

605

15

19.4

9.8

17.2

3.8

772

28

17.8

10.7

19.8

4.1

227

21.9

10.6

17.4

3.8

984

June

July

August

Sept*

Average

Least Significant Difference a t : 5%

5.6

1.7

2.6

.8

l%

7.6

2.4

3.6

1 .1

3^4. 468,

84

clipped frequently* came in late August*

Til© third#

and relatively minor peak#

No positive explanation can be of­

fered but it m a y have been caused by certain of the other species reaching maturity at that time* Note the similarity in the action of lignin to that Just described for dry matter*

There are minor inconsis­

tencies but the relationship is evident*

These relations

will be thoroughly discussed later* There is an average lignin content of 10*6 percent* This is higher than that found for either birdsfoot trefoil or Alta fescue alone, and demonstrates the fallacy of assum­ ing that the mixture is the result of the additive effect of its principal grass and legume*

The mixture is defin­

itely Influenced by the other species*

However# since no

chemical analyses were made of this fraction# its effect cannot be accurately determined* The average crude protein content of this mixture was 17*4 percent*

There is a progression from an initial high

of 2 0 * 2 percent in the early spring, to a seasonal low of 1 3 , 7 percent on June 22*

This is followed by a slow in­

crease to a second highest value of 1 9 * 8 percent on the last sampling date*

There is more than a sufficiency of

protein for beef animals even at its lowest level for the season* cent.

Ether extract had a seasonal average of 3*8 per­ In general, its fluctuations coincide with, but are

in a reciprocal direction to those described for lignin and dry matter*

Table 19 gives a summary by 6-week periods of the more detailed data presented in Tables 16, 17* and 13* Minor variations, which necessarily arise when sampling periods are only 2 weeks apart,

are reduced,

picture of seasonal trends, presented*

and a clearer

Table 19 Chemical Composition of Birdsfoot Trefoil, Alta Fescue, and Birdsfoot Trefoil - Alta Fescue Mixture Average of Three Two-Week Periods Chemical Composl tion - Percent Dry Crude Ether Fraction Matter Lignin Protein Extract X

Periods

BIRDSFOOT TREFOIL 1 st 2nd 3 rd 4 th

6 6 6 6

weeks weeks weeks weeks

Average

1 8 .5

7.4

13.5

1 0 .2 1 0 .2 8 .1

1 6 .4

9.0

3-7.3 1 6 .1

27.3 21.7 22.7

4.2 3.8 3.6

2 4 .2

£ .0

-----

2 ^ .0

3.9

-----

2.9 2 .7

ALTA FESCUE 1 st 6 weeks 2nd 6 weeks 3rd 6 weeks 4 th 6 weeks

Average

2 6 .8 2 6 .2 2 1 .4

1 0 .3 1 3 -2

1 6 .0 1 2 .6

3-9.5

20 *4

4 .3

19*7

9.5 7.3

^0.5 30.5 5 -0 . 0

4 .0

4 8 .6

23*3

1 0 .1

17.1

3.5

4 0 .0

BIRDSFOOT TREFOIL - ALTA FESCUE MIXTURE 1 st 6 weeks 2nd 6 weeks 3rd 6 weeks 4 th 6 weeks

2 1 .6 2 3 .8 2 2 .2 2 0 .1

9 .4 1 0 .6 1 0 .6

1 7 .6

3 .3

11.7

1 8 .4 1 8 .3

4*6 5.0

-------------

Average

21.9

1 0 .6

17 .4

3.8

-----

3-5.3

tm

m

tm

87

Results of the Chemical Analysis of Forage from a Permanent Bluegrass Pasture Receiving 60 Pounds Pounds K 2O, and 60 Pounds of Nitrogen Annually

The botanical composition of this pasture was as follows: Kentucky bluegrass

6 9 * 6 percent

Others 3 0 *Jj. percent (Includes timothy, redtop meadow fescue, white clover, weeds, etc*) All pastures received 60 pounds P 2 O 5

30 pounds ^ 0 *

This pasture received in addition 60 pounds of nitrogen annually*

One-half of this was applied in the early spring

prior to the start of the grazing trials*

The remainder was

applied between the two sampling dates, June 22 and July 7* For all practical purposes there were no legumes pres­ ent In this pasture*

White clover, the only legume listed

in the botanical analysis, accounted for only 1*6 percent of the total forage.

Only a mixture sample was taken for

chemical analysis*

Permanent Bluegrass Mixture Table 20* percent.

The average dry matter content was 26*5

This is higher than that shown for the four pas­

ture mixtures previously considered.

A high of 35*3 per­

cent was reached in July, a low of 18.9 percent Is shown In the late fall*

Only on this latter occasion did the dry

88

Table 20 Chemical Composition of Forage from Permanent Bluegrass Pasture Receiving 60 Pounds P^Oc, 30 Pounds KpO, and 60 Pounds of Nitrogen Annually and the Total Digestible Nutrients Produced Per Pasture (Sham leal Composition - Per cent ~ Dry Crude Ether Fraction Matter Llgnln Protein Extract_____ X_____ T.D.M.

Date April

28

29-9

8.9-

25.1

4.9

99-8

1618

May

12

27.7

7.1

1 7 .6

9-5

9-3.6

2226

26

2 0 .6

10.9

1 3 .(4-

9-o

33.3

2102

9

3 0 .0

11.5

15.6

if. 8

3 8 .2

1035

22

34*6

13.9

li+« 6

h.7

33.2

952

7

35.3

1(4-.7

13.8

h*5

3 1 .8

1616

21

2 3 .2

10.3

2 3 .0

6.6

9-3.3

721

k

2 6 .1

1 0 .5

22.6

6.8

96.2

1521

18

29-9

13.9-

21.5

5.5

39.0

1036

1

30.9

1 5 .0

20.2

5.4

39.3

1099

15

20.6

1 2 .3

22.8

5.5

m-3.3

620

28

18.9

11.(4-

22.9

5.6

q-3-L

593

26.5

11.6

19.5

5.2

14.0 . 0

1261

June

July

Aujust

Sept •

Averag e

Least S i g n i f i c a n t

Difference$ a t :

5%

3.9

1.5

9.0

2.2

3.6

629

1%

5.2

2.1

12.2

3.0

h-c

857

89

matter content ever drop below 20.0 percent.

Note the ef­

fect of the second application of nitrogen which was made shortly before the July 7 harvest.

At that time the sea­

sonal high of 35*3 percent had been reached; between July 7 and July 21 there was a drop of 12.1 percent.

The nitrogen,

aided by a heavy rainfall which had occurred on July 5# was responsible for this succulent growth. last long, however,

The effect did not

and by September 1 the dry matter con­

tent rose to 30*9 percent.

The drop during the last two

periods was due to general climatic factors, not to the con­ tinued effect of the nitrogen. The initial lignin content was 8.i+ percent.

Except for

a slight decrease during the second 2-week period, there was a steady rise in lignin to a seasonal peak of l4+*7 percent on July 7»

During the following 11+ days there was a de­

cided drop; in the short interval the lignin content fell 4*1+ percent.

This is principally the result of the mid­

season nitrogen application.

The rain, previously referred

to, which occurred on July 5. was a supplementary factor. Following this mid-season low of 10*3 percent, the lignin content again increased reaching a seasonal high of 15*0 percent.

During the last two periods,

there was the usual

decline• Crude protein exhibited a seasonal high of in the first period of growth.

percent

It then fell to a low of

13J+ percent in only two periods and remained low until the

90

period f o l l o w i n g

the n i t r o g e n application.

e secon d ar y p e a k of 23.0 p e r c e n t point o n the

Pr otein r e a c h e d

at tn at time.

trend is th e reciprocal

of that

From this

snown for

lignin w i t h the p r o t e i n d e c l i n i n g for a few p er i o d s before new fall g r o w t h a g a i n c a u s e s

an increase.

The a v e r a g e c r ud e fat c ontent of this f or a ge was c on ­ si derably h i g h e r than individual

s p ec i es

that f o un d for an y o t h e r p a st u re or

studied.

This w a s due to

the e x c e p t i o n ­

ally h i g h v alues o b t a i n e d a ft e r the n i t r o g e n was applied. There was a n in crease f r o m Ip.5 per c e n t o n July 7 to 6.6 p e r ­ cent o n J u l y

21.

O n a p e r c e n t a g e basis,

crude fat r e p r e s e n t s nitrogen.

the m o s t

this

increase in

s i g n i f i ca n t effect of the

T h e t r e n d shown for f r a c t i o n X is s im i la r to

that of the fat o r p r o t e i n contents. n i t r o g e n is c l e a r l y 11.5 p e r c e n t i^O.O pe r ce n t*

shown;

in one p e ri o d.

A g a i n the e ffect of

the f r a c t i o n X content increa s ed The

average seasonal

value was

91

Results of the Chemical Analysis of Forage from a Permanent Bluegrass Pasture Receiving 60 Pounds P205 and 30 Pounds K 20 Annually

The botanical analysis of this pasture was as follows: Kentucky bluegrass

69*8 percent

Others 30*2 percent (Includes white clover, timothy, redtop, meadow fescue, weeds, etc*) The percentage o f the main species, Kentucky blue­ grass, is almost identical to that found in the other per­ manent bluegrass pasture*

Other species, nowever,

contained

8.5 percent white clover as compared to the 1*6 percent in the previous pasture*

The 30 pounds of nitrogen added to

the previous pasture probably accounted for the scarcity of clover in that pasture*

This represents the only differ­

ence in the treatment of these two pastures.

P er m a n e n t B l u e g r a s s M i x t u r e

Table 21.

The average dry matter content, 29*9 per­

cent, is considerably higher than that of any other pasture mixture studied.

The difference between this value and

the 26.5 percent reported for the previous permanent pas­ ture is due to the nitrogen applied to the latter*

Note

the very high percentage of dry matter shown for the June 22 period*

Compare this with the almost identical value

reported in Table 13 for Kentucky bluegrass on this same

92

Table 21 Chemical Composition of Forage from Permanent Bluegrass Pasture Receiving 60 Pounds P 2 O 5 and 30 Pounds K P0 Annually and the Total Digestible Nutrients Produced Per Pasture Chemical Composition - Percent Dry Crude £ther Fraction Matter Lignin Protein iuctract X_____ T.D.M.

Date April

20

26.7

9.7

19.4

4.1

47.0

1031

May

12

21.0

8.8

15.4

3.9

46.6

1264

26

22.6

12.1*

13.9

3.5

36.7

1019

9

34.5

12.4

11.4

3.4

37.1

641

22

lfl.0

16.0

CVI •

-.53

-.0 1

-.35

.03 .11

Protein

73.07

37.87

CO EF'F1! CIENT OF DETiRKINATION - PERCENT (r? ) Lignin

1 2 .9 6

Dry Matter

9-00

.09

1 2 .96

5.29

I.69

9 .6 1

5.76

1 0 .89

Protein

#

Significance at the

5%

level - .57

Significance at the 1% level - .70

Table 30 The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil

Independent Vari ables

Dry Matter

Dependent V ari abl ss Crude fether Protein Extract

Fraction X

CORRELATION COEFFICIENT (r) .16

Lignin Dry Matter

-.82**

-•6?*

-.98*-

-.02

-.61*

-.23

Protein

•43

REGRESSION COEFFICIENT (by x ) .26

Lignin Dry Matter

-l.lj-8

-.12

-2.53

-.02

-.07

-.36

.05

1.20

Protein

COEFFICIENT OF D E T E R M I N A H O N - PERCENT (r2 ) Lignin Dry Matter

2.56

6 7 .?k •0i|.

Protein

3*.kk

96.04

37-21

5.29

18.^9

70.56

x

Prom Birdsfoot trefoil - bluegrass mixture

*

Significance at the 5 % level - .57

■a-* Significance at the 1 % level - .70

105

Table 31 Tbe Relationships Existing Between the Various Components Determined for Kentucky Bluegrass*

Independent Vari able s

£>ry Matter

Dependent Variables Crude Ether Protein Extract

CORRELATION COEFFICIENT (r)

-•Jtf

-.06

H •

1

Dry Matter

O in • 1

0J 1A •

Lignin

Protein

REGRESSION COEFFICIENT (by x ) 1.58

-.01

9

-.04

1

Dry Matter

-.91 0

Lignin

.15

Protein

COEFFICIENT OF DETERMINATION - PERCENT (r2 ) Lignin

27 «o2f

Dry Matter Protein

25.00

1.69

24.01

.36 .64

x

Prom birdsfoot trefoil - Kentucky bluegrass pasture

*

Significance at the 5% level - *57 Significance at the 1% level - .70

106

Table 32 The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil - Kentucky Bluegrass Mixture

Independent Vari ables

Dry Matter

Dependent Variables Crude Ether Protein Extract

T.D.N.

CORRELATION COEFFICIENT (r) •59*

•93**

CVJ

Protein

.16 •

-•50

0

-.30 •

Dry Matter

— 53

1

Lignin

-•58*

REGRESSION COEFFICIENT (byx )

-.38

Protein

rt

Dry Matter

-1.01

• 1

1.48

Lignin

39.93

-.05

43.38

.22

-7 8 .I45

COEFFICIENT OF DETERMINATION - PERCENT (r2 ) Lignin Dry Matter

3lf.8l

28.09

9.00

2.56

25.00

9.00

17.64

86.149

33.64

Pro tein *

Significance at the 5% level - .57

** Significance at the 1% level - .70

106

Table 32 The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil - Kentucky Bluegrass Mixture

Independent Vari ables

Dry Matter

Dependent Variables Crude Ether Protein Extract

T*D.N.

CORRELATION COEFFICIENT (r)

Dry Matter

-.53

.16

-.50

0 « 1

.59*

O • 1

Lignin

.42

Protein

•93**

-.58*

REGRESSION CO EFFICIENT (byx> Lignin

1.48

Dry Matter

-1.01

-.13

39.93

-.38

-.05

43.38

.22

-78.145

Pro tein

COEF FICIENT OF DETERMINATION - PERCENT (r2 ) Lignin Dry Matter

34-81

28.09

9.00

2.56

25.00

9.00

17.64

86.49

33.64

Protein *

Significance at the 5% level - .57 Significance at the 1 % level - .70

Table 33 The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil*

Independent Vari ables

Dry Matter

Dependent Variables Crude Ether Protein Extract

CORRELATION CO EFFICIENT (r) Lignin

.08

Dry Matter

-.57*

-.69*

.29

.37

Protein

.74**

REGRESSION COEFFICIENT (by x ) •li*.

Lignin

-1.07

-.16

.30

.05

Dry Matter

.09

Protein

CO EFFICIENT OP’ DETERMINATION - PERCENT (r£) Lignin

.64

32.49

47.61

8.41

13.69

Dry Matter

54.76

Protein x

From birdsfoot trefoil - Alta fescue pasture

*

Significance at the 5 % level - .57

-a-tt Significance at the

level - .70

108

Table 34 The Relationships Existing Between the Various Components Determined for Alta Fescue

Independent Vari ables

Dry Matter

Dependent Variables Crude Ether Protein Extract

fraction X

CORRELATION COEFFICIENT (r) Lignin

.55

Dry Matter

-.94 **

-.83-**

-.94**

—.66*

-.60*

-•*9

Pro t e in

,89-a-*

•93**

REGRESSION COEFFICIENT

-1.30 -.66

Dry Matter

-.26

-2.77

H • f

.76

Lignin

(h i >

-1.05 1.88

.21

Protein

COEFFICIENT Or DETERMINATION - PERCENT (r2) Lignin Dry Matter

30.25

88.36

68.89

88.36

43.56

36.00

24.OI

86.49

79.21

Protein *

Significance at the 5 % level - .57

** Significance at the 1 % level - .70

Table 35 The Relationships Existing Between the Various Components Determined for Birdsfoot Trefoil - Alta Fescue Mixture

Independent V ari ables

Dry Matter

Dependent Variable s Crude Ether Extract Protein

T. D. N.

CORRELATION COEFFICIENT (r) Lignin

.17

Dry Matter

-.09

• 13

-.51

-.0 6

• 37

-.30 -.1 2

Fro tein

REGRESSION COEFFICIENT (by*) .2 2

Lignin Dry Matter

-.1 0

.05

-IOI4..4O

-.0 6

.11

58.14

.23

-2 0 . 8 8

Protein

COEFFICIENT O r DETcJIMINATIOm - p rRCENT i s b Lignin Dry Matter

2 .89

.81

I.6 9

2 6 .01

.36

9.00

1 7 .6 4

Protein *

Significance at the 5/6 level - .57 Significance at the 1% level - .70

86.49

3 3 .64.

Tne 2

Ini^e e c i e c ’. L T i i r l f s

c e l ::on.

l o t i s : in^; I » for B l u e g r &ss M i xt v r — -

-^7

Ma:t«r

zr.L r.

Ztj

a

-reT^-an a *\

-a '

•» r . *

_ j

Leeender.c V i r i at 1 es Crude atder Ft i: 1 1 ;r Protein iXtri:: X

, .

Li

an

an. t s

,

.„C

• •

itd::er

* •

^

—m , ,

-

“ •

- • ~v.

* ~v

«'

•"*0

?r::eia

v- - - —*'~.'v -" a"_\ " ^"Wj X v

r e p r e s s : >.1

-1 #0 2

LlialS

-. r ?

2rj 5ia:ter

COEFFICIENT :? L ETER.Y I Lignin Lry Matter Pro t ein

*

-.r0

10.:3

.02

lO.PO

.Oo

?re c e in

21.16

C S - rePv. z.**a

_ 2 '*

^r

.Si

.Olf

SS.ifif

19.30

20.25

7 . 29

29.lt?

2 .to

is.;f9

50.25

I4 .OO

16

Significance at the 5~« level - .57

■> Significance at tne 1,^5 level - .70

,

^,

*-

-e

P. P. *

Ill

Iable 37 Iiie He 1 atloasnips Existing Between the Various Caapoaeot s Determined for Permanent Bluegrass Mixture - FS

Independent V?-!ables

Dry Matter

Dependent Variab les Crude Ether Fracti on Protein Extract X

T. 0. X

CO HR.EEATI 0 i» COEFFICIENT (r) 1i»ni r.

•71**

-.2= — m5-5^

_ry Matter Protein

.23

-.71*-*

— .00

-.77*-*

— •-*-0

.5^

-.Id

*00

-.64

-17.33

.13

1.26

-17.20

•5**

REO-REBSIOH CO Er r I 01EN i. (by^ ) 1.34

Liynin -ry Matter

-.30 -.27

Protein

COEFFICIENT Or DETERMINATION - PERCENT (r2 ) Lignin Cry Matter Protein *

50.41

6.25

5.29

73.96

50.41

33.64

.00

59.29

16.00

29 .1-6

29.10

3.24

Significance at tne 5l* level - .57

*■* Significance at the 1 % level - .70

11

Table 3® la * compilation of the correlation coeffi­ cients found for each species and mixture*

Tne & roupia^

of this material facilitates comparisons and the summariza­ tion of the relationships noted* Considering all species and mixtures involved,

there

appears to be a rather definite positive correlation be­ tween lignin and dry matter*

Tne smaller coefficients

shown for the legumes would indicate that they are less likely to show such a relationship than are the grasses or tne mixtures.

This can be reasonably explained on the

bases of tne lower lignir and dry matter contents of the legumes and the smaller deviation of tne periodic values of tnese constituents around their seasonal mean.

Although

there were only three statistically significant correlation coefficients attesting to this relationship,

the consistency

of the trend would indicate considerable reliability. Lignin appears to be negatively correlated with the crude protein content of the plant. dicate a stronger,

Larger coefficients in­

more consistent relationship between

these two variables than the two previously discussed. Seven correlation coefficients out of fourteen calculated were significant or highly significant* closely approached these odds*

Many of the others

This relationship appears

to be stronger for the legumes and grasses then for the mixtures.

Tnia may be due to tne effect of the other

species included in the mixture samples.

113 v>1 - J*wi.

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1114-

Lignin extract

shows

and f r a c t i o n X.

ere s i g n i f i c a n t tion.

& negative

but

A fcain the

there is

evidence

such rel ationsni ps :dxtures.

Not

all o f trie c o e f f i c i e n t s found a d ef i n i t e

tne p e rc e n t panied by

in the g r a s s e s

tnat w n e n

a decrease

the e x i s t e n c e of

and l e g u m e s

tn an for the

this

i nc r ea s es

in t u r n is a c c o m ­ protein,

and f r a c t i o n X fo u nd in tne p l a n t

gestible n u t r i e n t s

p r o d u c e d p e r pasture

T h re e o f tne m i x t u r e s

broriegrass,

birdsfoot

but

e v e r a t ed,

ignored.

was

a t re n d

Ladino a nd

in d iv i d u a l

significant•

Any positive relationship

d ep r es s in g

effect

c lover -

coefficient

i n d i c a t e d v.’h i c h cannot

is in direct co n tr a st

to

c o mb i ne d c o e f f i c i e n t when

found to be

and T.u.N.

di­

alfalfa -

correlation

Neither tne tnree

c o r r e l a t i o n c o e f f i c i e n t s n o r tne

h o we v e r ,

studied,

positive

tnese t w o v a r i a b l e s .

There is,

are d i f f i c u l t

t r e f o i l - bluegress,

s n ow e d e .wall

were

tissue.

found b e t w e e n l i g n i n a n d total

surnr.ari ze.

the three

ere not there are

the l ignin c o nt e n t

in trie co ntents of cr u de

The r e l a t i o n s h i p s

pletely

in tni s d i r e c ­

in eacn case studie d,

dry nfctter increases;

et.ner e x t r a c t

between

trend

A 1 t h o u g h the c o r r e l a t i o n c o e f f i c i e n t s

indications

timothy,

to b o t n ether

is s t r o n g e r for

statistically significant strong

relationsnip

be c o m ­

i n v o l v i n g l ig n in

to p r es e nt k n o w l e d g e of tne

of an i n c r e a s e d l ignin co n t e n t on the d i ­

g e s t i b i l i t y of forages. tion o f this unusual

It is b e l i e v e d that

s i tu a t i o n can be found

tne p o s i t i v e r e l a t i o n s n i p

snown b etween

tne e x p l a n a ­ by c o n s i d e ri n g

dry m a t t e r and

115

T.D.N.

Since lignin is positively related to dry matter,

end dry matter is positively correlated with T.D.N., lignin would necessarily be positively related to T.u.N.

It would

appear that dry matter rather than lignin was the limiting factor in the production of T.D.N. on these three pastures. Tne percentage of legunes is higher in these pastures than cn the three remaining pastures which did not show this contradictory relationship or trend. shown In the discussion of tne

It has already been

chemical composition of the

forage that the legumes are extremely certain periods.

low In dry matter at

During these periods the animals may not

have been able to consume sufficient green forage to meet their daily dry matter requirements.

The laxative effect of

the lush fora6e may have been a n additional factor* Three of the pastures, birdsfoot trefoil - Alta fescue, permanent bluegress - N-P-K, eund permanent bluegrass - P-K, snowed tne expected negative r©lationships between lignin and T.D.N.

As tne lignin increased the T. j-'.N. produced per

pasture decreased,

and visa verse.

Although only one of

the three individual correlation coefficients was signifi­ cant, the coefficient obtained, by averaging the three was found to be highly significant. are predominately grass.

Tnis

Note that these pastures is especially true for the

two permanent pastures which also snow nigh dry matter and lignin contents. The percent dry matter generally showed an inverse

r e l a t i o n s h i p w i t h pr o t e i n , M an y of

tnese

correlation

ally s i g n i f i c a n t

coefficients

and in several

r e l a t i o n s h i p w a s noted. No e x p l a n a t i o n

e t h e r extract,

timothy,

were

g es t ur e s

dry m a t t e r

It is

None

content

Dry matter

and al f a l f a -

c ou l d h a v e

to note

tne l i g n i n

how­

these

p r o p o r t i o n of succulent

b e e n a limi ting

factor.

were p r e d o m i n a t e l y grass.

that these

show low but n e ga t i v e

c o r r e l a t i o n s b e t w e e n dry m a t t e r and T.D.N. e x p e c t e d si n ce

reported,

As p o i n t e d out p reviously,

three remaining pastures

interesting

clover - bromegrass,

of the c o e f f i c i e n t s

significant.

inconsistencies.

a p p e a r e d to be p o s i t i v e l y

contained a considerable

legumes. The

La d l n o

- X e n t u c k y bluegrass,

r el a t e d to T. D . N* ever,

i s o l a t e d cases a p o s i t i v e

is o f f e r e d fo r t h es e app arent

trefoil the

w er e not s t a t i s t i c ­

N o ne of t hese were signi f ic a nt *

For the t h r e e pastures, birdsfoot

and f r a c t i o n X.

This would be

c ontent of t he s e p a s t u r e s

a pp eared

tc be p o s i t i v e l y c o r r e l a t e d wi tn dry m a t t e r and n e g a t i v e l y correlated T wo of

to T.D.N. the most

consistent

associations

found were b e ­

tween p r o t e i n and e t h e r e xt r ac t or f r a c t i o n X. vidual

c o rr e la t io n s,

highly

s i g n i fi c an t ,

t iv e ly re l a t e d .

m o s t of w h i c h were i n d i c a t e d that

This

case of f r a c t i o n X,

would be expected,

since a c o n s i d e r a b l e

s u b s t a n c e is p r o t e i n . shown

these

The i n d i ­

s i g n i f i c a n t or f a ctors wer e p o s i ­ e s p e c i a l l y in the po r ti o n of this

There is a n e g a t i v e r e l a t i o n s h i p

in the m i x t u r e s b e t w e e n p r o t e i n and T.D.N.

production.

117

A l th o ug h o n l y culated w e r e

two of

the six c o r r e l a t i o n

signif ic a nt ,

t ne

sense

ship s u b s t a n t i a t e s not a l i m i t i n g There

are

of ca u se

Tnis

and ef fect.

f a c t o r at any time

cannot.

cannot be m e a s u r e d by yearns d a t a w i ll snould f a c i l i t a t e

in­

This relation­

during the g r a z i n g

certain inconsistencies

others

s n o u l d n o t be

the g e n e r a l c o n c l u s i o n that p r o t e i n was

large g r o u p o f c o r r e l a t i o n s . plained,

cal­

i n d i c a t i o n of this i n verse

relati on w o u l d a p p e a r r e l i a b l e * t er p r et e d in tne

coefficients

tne

Som e

present

season*

in this

can be l o g i c a l l y e x ­

y a n y i n t e r r e l a t i o n s exist wh i ch s t a t i s t i c s em p lo y ed .

A se c o n d

be o b t a i n e d o n m u c h o f this m a t e r i a l

and

t n e c l a r i f i c a t i o n o f q u e s t i o n a b l e points.

S UMMARY AND CONCLUSIONS

A study was m a d e of the seasonal variation in the chemical composi ti on of six rotationally grazed pastures at the Miller-Purdue Farm during the 19if9 season* of these pastures were grass and legume mixtures,

Four two were

unploughed permanent pastures consisting almost wholly of grans* This investigation was conducted in conjunction with grazing trials.

From these trials it was possible to ob ­

tain the infonna ti o n necessary to calculate the total d i ­ gestible nutrients pro d uc ed per pasture for each lip-day period during the grazing season*

These periods corre­

sponded to the periods at w h i ch the forage was sampled for chemical analysis* The percentage of dry matter found in all forages was unusually h igh in the early spring. came in early or mid-summer,

The peak generally

after which there was a de­

cline to a seasonal low in the fall.

The dry matter con­

tent varied widely bet we en species and between mixtures* Legumes ranked lowest in dry matter content, with Ladlno clover showing a seasonal average of 16.3 percent.

The

lowest individual value was 8.2 percent reported for birdsfoot trefoil in the late fall*

The average dry matter

content of Kentucky bluegrass was 31*2 percent, than any other species or mixture.

higher

119

The lignin content was lowest in the early spring growth,

increasing to a seasonal peak at the time the Tor-

age reached maturity, in the fall*

and then decreasing to a lower value

This later decline was due to the resumption

of vegetative growth*

The legunes were generally lower

than the grasses with Ladlno clover showing the lowest sea­ sonal average*

The variation of the periodic values around

the mean was relatively small,

Indicating very little change

in the lignin content during the season* age lignin value, bluegrass*

The highest aver­

13*5 percent, was reported for Kentucky

This Is considerably higher than the 10*1 per­

cent found for Alta fescue for which a h igh value might have been predicted on the basis of its physical character­ istics* The crude protein content was highest in the spring and fall, declining to a seasonal low during the early or mid-summer periods*

Birdsfoot trefoil showed the highest

seasonal average but was closely followed by the other legumes*

The highest individual value, 32»0 percent, was

reported for Ladlno clover*

At no time did the protein

content of any species or mixture drop below the minimum protein requirements for maximum beef production* The ether extract was highest in the spring and fall* Tne intermediate values were tion*

subject to considerable varia­

On the basis of tne small amount found,

and the low

digestibility reported for this heterogeneous substance in

120

the literature,

it is n o t believed to be of much nutritional

significance* The trends shown by fraction X closely parallel those previously described for crude protein* ages were m u c h higher*

The actual percent­

The highest content of fraction X

was found in the legumes* The relationships between the chemical constituents were determined for all species and mixtures* were also determined between the lignin,

Relationships

dry matter and

crude protein contents and tne total digestible nutrients produced per pasture for each mixture studied* There seemed to be a general positive relationship be­ tween lignin and dry m a t t e r content.

Lignin was found to be

inversely related to crude protein, ether extract and frac­ tion X*

Not all of the individual correlation coefficients

were statistically significant but a trend was indicated* Lignin appeared to be positively correlated with T*D.N* on the three pastures which contained relatively large amounts o f succulent legumes*

This was due to the associa­

tive effect of lignin and T.D.N* with the dry matter con­ tent*

This suggests that for pasture mixtures containing

a high amount o f succulent legumes,

the percentage dry

matter rather than the lignin content m a y be the limiting factor in beef production* Three pastures which were predominately grass showed an inverse relationship between lignin and T.D.N*

It would

appear that the lignin content is an important factor af­ fecting the nutritive value of forage which has a relatively high lignin and dry matter content* An inverse relationship was Indicated between dry matter and crude protein, ether extract, or fraction X* When the dry matter content was relatively low, dry matter appeared to be positively related to T.D.N.

The reverse was

true for the mixtures which had high dry matter contents* More data are necessary before drawing definite conclusions as to the relationships between lignin or dry matter and T.D.N. production* A mid-season application of nitrogen to a permanent bluegrass pasture resulted in a decided decrease in the lignin content of the forage.

The lignin content of the

permanent bluegrass pasture which did not receive this treatment continued high throughout the season*

122

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2.

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a.

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APPENDIX

s p•H
ac • 0

• CM xO

pH

Sept

(CM I dO •H d UN u 1 •H 0 pH 1

UN • O ' UN

m

July

cm

1 ■d rH0 ® 44 O UN 1 « • d pH p ■ u < 0

P*-

June

4* cm O 1 m0 u +» * u> w 1 CQ u • H A 4> • «
+» O 4-> o LA U 1 CM CQ o rH

in pounds

E-t

(h

r—

rH

vO

is reported

a

O' XA CM

T.D.N, which

iA

5"

A

Excepting

k\j ■ o

Date

Detailed

Data on the Chemical

Composition

of I.adino

Clover

- Bromegrass

Mixture

- Percent#

133

131*

( 4A ■%

1

.c? Ci

1

Q « -P a. O

CM

I o

CO

A a

•ft LA c 1 ■H

on the Data

1

O

CJ A



A

-P -P aj LA as 1 >> Q

0 CM

I CQ

r— A

O ' • CM

CM • A

-d * A

ao • A

nO

• CM

rH • A

O • A

O ' • CM

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ao • A

^0 •

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A • A

A

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rH

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A

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rH • A

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co

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A

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• CO

• A

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1

Date

Chemical

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LA

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-



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m

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A

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8

CM

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X

• CM

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-d* • CA

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