Volume 12, Number 6 (November 1959)

In This Issue

Growth and Carbohydrate Trends in Crested Wheafgrass D. N. Hyder and F. A. Sneva

Effects of Presowing Vernalization on Survival and Developrl?ent of Several Grasses...NeiZ C. Frischknecht

A Rancher’s ideas on Range Capacity Determinations

Dan G. Freed

The Effecf of Site on the Palatability and Nutritive

Conienf of Seeded Wheafgrasses...C. Wayne Cook Geomorphology of fhe Southern Great Plains in

Relation to Livesiock Producfion...WakefieZd Dort, Jr.

A Pasture Comparison Method of Estimating Utilization of Range Herbage on the Central Great Plains

R. E. Bement and G. E. Klipple

Germinative Characteristics of Grass Seed under

Snow... __._ ____________ .____ ___________ _____._______________ A. T. Bleak

Volcano Ranching: Problems and Opportunities in

Management of Hawaiian Range Land ____________ E. J. Britten

A Comparison of Two Grass Sampling Methods for Digestibility Trials Conducted on Pasture

E. F. Smit,h, V. A. Young, L. A. Holland, and H. C. Fryer

Changes in Interspecific Associations Related fo

Grazing Pressures..._...___._~~._~.__~Di~ie R. Smith

Book Reviews: Environmental Conservation (Dasmann) ;

Nomenclature of Plants (St. John); The Sheep

Book (McKinney); Fundamentals of Ecology (Odum)...

Current Liferafure... _______ ____ ____ __ _________ ________..___ _______.__ .-Lee A. Sharp

Society Business: Teniative Program 13th Annual

Meeting: Report of Professional Standards Commiffee... With the Sections... ____ _____________________ ____ _______ __________.._ ___ _________ ____ News and Notes ____ ______ _..__._ _______.______________ ____ _______________ ______ __________ ____ ___ ____ _

271

280

287

289

296 298 303

306

309

312 315

American

Society of Range Management

The American Society of Range Management was created allied technologists, and to encourage professional improvement in 1947 to foster advancement in the science and art of grazing of its members.

land management, to promote progress in the conservation and Persons shall be eligible for membership who are interested greatest sustained use of forage and soil resources, to stimu- in or engaged in practicing range or pasture management or late discussion and understanding of scientific and practical animal husbandry; administering grazing lands; or teaching, range and pasture problems, to provide a medium for the Dr conducting research, or engaged in extension activities in exchange of ideas and facts among society members and with range or pasture management or related subjects.

T

Off ice, Executive Secretary. Address all inquiries and correspondence including memberships, renewals, replacements of JOURNALS, etc., to Executive Secretary, American Society of Range Management, P.O. Box 5041, Portland 13, Oregon.

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Reprinting. The reprinting of articles or parts of articles published in the JOURNAL OF RANGE MANAGEMENT is author- ized on the express condition that full credit be given the JOURNAL and the author. The date of original publication must be shown with the credit line.

Posf Office Enfry. Second-class post- age paid at Portland, Oregon, and at ad- ditional offices.

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Printers. The Nebraska Farmer Company, 1420 P Street, Lincoln, Nebraska.

Copyright 1959 by the American Society of

Range Management.

OFFICERS OF THE SOCIETY President:

DONALD F. HERVEY Colorado State University

Fort Collins, Colorado

Vice President: Executive Secretary:

FRED H. KENNEDY JOHN G. CLOUSTON

U. S. Forest Service P. 0. Box 5041

Albuquerque, N. Mex. Portland 13, Oregon

BOARD OF DIRECTORS 1957-59

KENNETH CONRAD E. WM. ANDERSON

Wray, Colorado Soil Conservation Service

Pendleton, Oregon

1958-60

JAMES L. FINLEY MELVIN S. MORRIS

Holbrook, Arizona University of Montana

Missoula, Montana 1959-61

JOHN CHOHLIS C. H. MCKINNON

Western Livestock Journal

Sacramento, California LK Ranches, Ltd. Calgary, Alberta

Past President: ROBERT S. CAMPBELL

U. S. Forest Service New Orleans 13, La.

HISTORIAN: A. A. BEETLE, Dept. of Agronomy, University of Wyoming, Laramie, Wyoming

JOURNAL OF RANGE MANAGEMENT EDITOR

WARREN C. WHITMAN

Department of Botany, North Dakota Agricultural College Fargo, North Dakota

EDITORIAL BOARD

1957-59

FRANK GYBERG E. J. WOOLFOLK

Cornville, Arizona Pacific Southwest Forest & Range Exp. Sta. Berkeley 1, California 1958-60

ARNOLD HEERWAGEN W. R. HANSON

Soil Conservation Service

321 New Customs Building Eastern Rockies Forest Denver 2, Colorado Conservation Board Calgary, Alberta

1959-61

DONALD R. CORNELIUS JACK R. HARLAN

Agricultural Research Service Oklahoma State Universitl

P. 0. Box 24’5 StillwPtcr. Oklahoma

Journal of

RANGE

MANAGEMENT

Growth and Carbohydrate Trends in Crested

Wheatgrass’

D. N. HYDER AND F. A. SNEVA

Range Conservationists, Crops Research Division, Agri- cultural Research Service, U. S. Department of Agricul- ture, Burns, Oregon

The importance of carbohy-

drate reserves in perennial plants

is well recognized and docu-

mented. The seasonal trend in

depletion of reserves for initial

growth and subsequent replen-

ishment has become an impor-

tant consideration in range man-

agement due to the fine contri-

butions of Sampson and Mc-

Carty (1930)) McCarty and Price

( 1942)

McIlvanie (1942) ,

Wein-

mann (1952)) and many others.

This paper presents growing-

season trends in herbage produc-

tion and root carbohydrates by

crested wheatgrass

desertorum).

Procedure

The work reported was con-

ducted on Squaw Butte Range

in southeastern Oregon. The

bottom sites chosen for study

represent the deeper, relatively

rock-free soils in this area. Blue-

bunch wheatgrass

X’ontribution from Squaw Butte- Harney Experiment Station, Burns, Oregon. This station is jointly oper- ated and financed by the Crops Re- search Division, Agricultural Re- search Service, U.S.D.A., and Oregon Agricultural Experiment St at i o n, Corvallis, Oregon. Technical Paper No. 1202, Oregon Agr. Exp. Sta.

spicatum),

sandberg bluegrass

(Poa

and Junegrass

(Koeleria cristatu)

are dominant

native grasses found with big

sagebrush

on these soils. Average precipi-

tation is slightly over 11 inches.

In 1956 crested wheatgrass

plants were removed from the

field with a column of undis-

turbed soil 12 inches in diameter

and 18 inches deep for potting.

These were mature plants that

had been seeded in 1952 and sub-

sequently protected from graz-

ing. Thirty-two plants were

potted and arranged in a plastic-

lined trench. that was refilled

with soil to maintain field con-

ditions with respect to soil tem-

peratures. Potting was completed

March 20-27, 1956-about 10 days

after snow melt. The plants

were arranged by clump sizes

into 4 groups identified as A

(large)

and D (small).

Each pot was fertilized with am-

monium nitrate at a rate equi-

valent to 20 lb. N/A. Applica-

tions were made in 1 quart of

water. Additional irrigation of

1 quart was provided on June 12.

One pot of each size class was

removed on each of 8 dates at 2-

week intervals beginning April

24 and concluding July 24. The

herbage was clipped from the 4

271

pots at ground level, dried in a

forced-air electric oven at 9O”C.,

and weighed to the nearest gram.

Stem bases and roots were wash-

ed from the soil column, dried

at 9O”C., weighed, ground in a

Wiley mill to pass through a 20-

mesh screen, cornposited, and

preserved in sealed containers

for chemical determinations of

total carbohydrates.

In 1957 studies were conducted

under field conditions on 2 simi-

lar sites identified as site I and

site II. On site I, 4 randomized

blocks of seven 48-square-foot

plots each were prepared and

cleaned of old herbage in the fall

1956. On this site there was a

mature stand of crested wheat-

grass that had been drilled in

la-inch rows in the fall 1952 and

subsequently protected

from

grazing. Herbage was harvested

by hand clipping near ground

level on 1 of 7 dates at a-week

intervals beginning May 7, 1957

and concluding July 30. The

herbage was dried at 90°C. and

weighed to the nearest gram.

Also, root samples were taken

from clipped plots on each date.

Root samples, including 5 or 6

clumps on each plot, were ob-

tained by excavating to a depth

of about 8 inches at a distance

of 6 inches from the crowns. The

roots with stem bases were

washed from the sods, dried,

ground in a Wiley mill, compo-

sited, and sealed in glass con-

tainers for chemical determina-

tions.

272

D. N. HYDER AND F. A. SNEVA

Table 1. Precipifafion and atmospheric temperatures fhaf provided favor- able growing conditions in 1956 and 1957.

Monthly precipitation (inches) Mean monthly temp. (OF.1 Month ‘55-56 ‘56-57 ZO-yr. mean *‘55-56 ‘56-57 20-yr. mean

October 0.49 2.57 1.01 49 48 48

November 1.80 0.18 1.11 34 38 36

December 2.05 0.75 1.46 31 31 29

January 3.20 1.40 1.24 29 18 25

February 1.00 1.19 1.03 26 34 29

March 0.65 3.21 0.82 36 38 34

April 0.20 0.75 0.67 47 44 43

May 3.04 2.70 1.40 52 51 50

June 0.80 0.24 1.34 55 59 56

Totals 13.23 12.99 10.08 359 361 350

out treatment for about 15 years.

Individual plots were 8 x 12 feet

in size. The 2-level factor in-

cluded untreated and deheaded

plants. Deheading was accomp-

lished by cutting the culms at

the base of the heads shortly

after they emerged from the

boot. The la-level factor includ-

ed 12 dates of harvest at weekly

intervals beginning .June 5, 1957

and concluding August 21. Herb-

age and root samples were taken

on each date and processed in

the manner described.

Carbohydrate concentrations

were determined by the Shaffer

and Hartmann modification of

the Munson and Walker method

(Browne and Zerban, 1941) .2

This determination obtains most

of the carbohydrates in forms

other than cellulose and lignin.

The results are presented as

total-carbohydrate

concentra-

tions in terms of glucose equi-

valent.

The growing seasons in. 1956

and 1957 were unusually favor-

able. Precipitation and temper-

ature records are presented in

Table 1.

Resulfs

Crested Wheafgrass Grown In Pofs

Growing-season trends in the

total-carbohydrate

content

of

Whemical determinations were per-

formed by the Dept. of Agricultural Chemistry, Oregon State College, Corvallis, Oregon.

roots with stem bases and in

herbage dry-matter yields by

crested wheatgrass grown in

pots during 1956 are presented

in Figure 1.

Carbohydrate determinations

were performed on composite

samples. Mean values of tripli-

cate determinations are pre-

sented.

Analyses of herbage yields

gave highly signif icant differ-

ences among clump-size classes

and among dates of removal.

Mean herbage weights were 14,

13, 11, and 10 grams of dry mat-

ter, respectively, by size classes

A (large), B, C, and D (small).

Consequently, stratification by

size classes was an important ex-

perimental control. Among dates

of removal the final significant

increase in yield was obtained

in the period June al-July 2, as

measured by the 5 percent L.S.D.

of 3.7 grams per pot. Mean herb-

age weights of plants were 19,

18, 16, and 16 grams, respective-

ly, by size classes A, B, C, and

D when clipped on and after

July 2.

Analysis of root yields (with

stem bases) gave highly signifi-

cant differences among

size

classes, but dates of removal did

not give significant variation. In

fact, root weights did not mani-

fest even a slight tendency for

seasonal increases. Clump size

stratification was of importance

in root weight. Mean root weights

were 66, 60, 53, and 37 grams of

dry matter, respectively, by size

classes A, B, C, and D. The root:

top ratio was 3.1 to 1, as com-

puted from the mean root weight

of all plants and the mean herb-

age weight of plants removed on

and after July 2.

The data of Figure 1 suggest

that: (a) The depletion of car-

bohydrate reserves for initial

growth probably was not ex-

cessive, (b) the accumulation of

carbohydrates proceeded rapidly

during the growing season and

by head emergence in early June

was high at about 27 percent,

(c) herbage growth was com-

pleted by flowering time, and

(d) the termination in herbage

growth and initiation of repro-

ductive activity were associated

with a moderate decrease in car-

bohydrates that was not restored

until after the seeds were filled.

Crested Wheafgrass Grown in The Field

Growing-season trends in the total-carbohydrate content of roots (with stem bases) and herbage dry-matter yields by crested wheatgrass grown in the field during 1957 are presented

in Figure 2.

Carbohydrate determinations

were performed on composite

samples. Mean values of tripli-

cate determinations are pre-

sented.

Statistical analysis of herbage

yields gave highly signif icant

differences among dates of har-

vest. The final significant in-

crease in yield was obtained in

the period June 4-18, as meas-

ured by the 5 percent L.S.D. of

330 lb/A. However, yield sam-

ples taken on and after July 1

averaged 1,490 lb/A, or 150 lb/A

higher than on June 18.

CARBOHYDRATES IN CRESTED WHEATGRASS

273

>o,

I Oo\o

m-

a

2

24 APRIL

8 21 4 21 2

MAY JUNE

REMOVAL DATE (1956)

17

JULY

24

FIGURE 1. Seasonal trends of total carbohydrates in roots (with stem bases) herbage yields by crested wheatgrass grown in pots.

a slight decrease in total carbo-

hydrates was associated with or

slightly preceded the termina-

tion in herbage growth and initi-

ation of flowering, and (d) a

moderate increase in total carbo-

hydrates occurred during the pe-

riod of seed formation.

Figure 2 data began too late in

the season to provide informa-

tion pertaining to the level of

carbohydrate depletion with ini-

tial growth. On May 7, when

herbage production was just 24

percent complete, total-carbohy-

drate concentrations were over

23 percent.

Effects of Deheading upon Carbohydrate trends

Pre- and post-flowering trends

in the total-carbohydrate content

of roots and stem bases by un-

treated and deheaded crested

wheatgrass plants grown in the

field during 1957 are presented

in Figure 3. Corresponding herb-

age dry-matter yields are pre-

sented in Table 2.

A paired comparison analysis

of root carbohydrate values indi-

cates no differences due to de-

heading. The mean difference

and of

was 0.4 percent glucose equiva-

lent, and the standard deviation

of individual differences was

1.23 percent. Since the greatest

difference was just 2.2 percent

on June 26, it seems likely that

deheading did not influence root

32

1

carbohydrate levels at any time.

The primary objective in de-

heading was to find whether the

carbohydrate decrease in June

was directly associated with

flowering activity. Root carbo-

hydrates decreased

in both

treatments, although the de-

crease was slightly less in de-

headed plants.

Statistical analysis of herbage

yields gave highly signif icant

differences among the 4 replica-

tions and among the 12 dates of

removal.

The differences

in

yields by untreated

and de-

headed plants were significant

at the 5 percent level. However,

deheading removed an average

of 53 lb/A. of dry matter. Since

the 5 percent L.S.D. for testing

treatment means was 80 lb/A,

it is clear that the removal of

heads was responsible for the

significant difference. No fur-

ther effects of deheading were

apparent. Among dates of har-

vest the final increase in yield

occurred in the week June 26-

July 3.

Considering mean carbohy-

drate values and herbage yields,

r

28,

-800

- Root carbohydrates ---- Herbage yields

-600

,C N *& -400

22,

Seeds -200 mature

Of 0

I 1 I I I

7 21 4 I8 I 16 30

MAY JUNE JULY

REMOVAL DATE (1957)

274

D. N. HYDER AND F. A. SNEVA

the data presented in Figure 3

and Table 2 suggest that: (a)

Total carbohydrate levels were

about 27 percent at heading time

in early June, (b) herbage

growth terminated by, or dur-

ing, flowering activity, (c) a

moderate decrease in total car-

bohydrate occurred by June 26

when flowering activity

was

just beginning, but recovery

was prompt in the subsequent

week during greatest flowering

activity, and (d) total-carbohy-

drate levels fluctuated between

25.5 and 30 percent throughout

July and August.

By August 21 the grasses were

brown, and the herbage con-

tained only 34 percent moisture.

It seems unlikely that root stor-

age levels increased at later

dates.

Discussion and Conclusions

Growing-season trends in the

total-carbohydrate

content

of

crested wheatgrass

roots (with

stem bases) appear to be char-

acterized by:

(a)

An early

rapid increase during the grow-

ing season to a level near 27 per-

cent at about the time of head

emergence, (b) a moderate de-

crease at, or just prior to, flow-

ering, and (c) a final recovery

during or just following seed

formation to a level approxi-

mately the same as that attained

by head emergence.

An important period for the

accumulation of carbohydrates

coincided with the growing sea-

son. Reserves were restored

most rapidly in May prior to

head emergence in early June.

The time and manner of deple-

tiDn of reserves

for initial

growth w a s not investigated.

However, it is recognized that

the extent of depletion for initial

growth might strongly affect the

rapidity and time of recovery.

Some concern regarding

the

conclusion of early accumulation

of carbohydrates

in crested

wheatgrass must be expressed,

because delayed accumulation is

sometimes represented as the

Z -

Seeds

Oeheading Flowering mature

I I

5

12 19 26

3

I

IO 17

I

24

I

I

I I4 21 T

JUNE JULY AUGUST

REMOVAL

DATES

(1957)

FIGURE 3. Pre-and post-flowering trends of total carbohydrates in roots (with stem

bases) by untreated and deheaded crested wheatgrass plants grown in the field.

classic pattern regarding food

reserves.

Delayed accumulation of re-

serves may be illustrated with

data on California needlegrass

Total carbohy-

drates in the stem bases of Cali-

fornia needlegrass

were low

(about 4 percent) during the

season of active growth (Samp-

son and McCarty, 1930). Flower-

ing occurred

when herbage

growth was about 40 percent

complete. Thereafter the com-

posite demands of growth and

reproductive

activities

con-

sumed current production of car-

bohydrates. Terminal

storage

levels obtained after termination

in growth and reproductive ac-

tivities were about 17 percent.

In contrast with California

needlegrass, flowering and seed

formation are delayed in many

species until herbage growth is

essentially complete.

Neiland

and Curtis (1956) listed the dur-

ation of pre-flowering vegetative

growth as an important factor in

the ability of a grass to with-

stand grazing. It seems likely

that the accumulation of food

reserves is related to the se-

quence in growth and repro-

ductive activities.

Moderately early accumula-

tion of carbohydrate reserves

may be illustrated with data re-

garding slender wheatgrass

ropyron trachycaulum),

which

CARBOHYDRATES IN CRESTED WHEATGRASS 275

portance in slender wheatgrass.

However, it is recognized that

there is sufficient opportunity

for controversy.

Early accumulation of carbo-

hydrates may be illustrated with

data regarding orchardgrass

(Dacty Zis glomerata). Concen- trations of food reserves (sugars

and fructosan) in the stubble

decreased to about 12 percent

shortly after clipping and in-

creased to about 26 percent in 35

days after clipping (Sullivan

and Sprague, 1953). Reserve

levels in the roots were lower than in the stubble, but followed a similar trend of increase from about 10 percent to 19 percent in 35 days after cutting.

A number of factors other

than the duration of pre-flower-

ing vegetative growth may con-

tribute to differences in the pat-

tern of carbohydrate accumula-

tion in grasses. Morphological

differences in the ‘origin of

photosynthetic tissue may be of

special interest in range manage- ment. Branson (1953) discussed the importance of vegetative: re-

productive stem ratio and the

height of the growing points.

The occurrence of basal leafiness in many species is easily recog- nized in the field. Species hav-

ing vegetative stems, in which

the internodes are very short

and produce only leaves, appear to have good structure for early

development of abundant photo-

synthetic tissue as well as for

the replacement of photosyn-

thetic tissue removed by grazing. Species having a high propor-

tion of reproductive stems ap-

pear to develop abundant photo-

synthetic tissue more slowly,

and to replace it with greater difficulty after grazing or clip- ping than those with vegetative

stems. In reproductive stems

short basal internodes may per-

mit more rapid development of

photosynthetic tissue than long

basal internodes, because a leaf

arises at each node. Also, as

emphasized by Branson (1953))

Table 2. Pre- and post-flowering trends in herbage dry-matter yields by untreated and deheaded crested wheatgrass plants grown in the field.

Removal date (1957)

Herbage dry-matter yields (lb/A) by plants: Untreated Deheaded” Average

June 5 701 626 6641_

June 12 902 862 882

June 19 829 786 808

June 26 970 1006 988

July 3 1316 976 1146

July 10 1057 952 1005

July 17 1268 981 1124

July 24 1160 1084 1122

Aug. 1 1160 1058 1109

Aug. 8 1076 1112 1094

Aug. 14 1074 924 1000

Aug. 21 966 994 980

Average -

*

short basal internodes delay the

rise of growth primordia to a

grazable height.

Early growth in crested

wheatgrass was characterized by

abundant leafiness, delay in

stem elongation, and opportun-

ity for the accumulation of car-

bohydrates. The abundant leafi-

ness at an early stage of growth was attributed to short basal in- ternodes, because each stem pro-

duced a reproductive culm.

Crested wheatgrass produced 2

or more short internodes below the soil surface as described by

Cook and Stoddart (1953). For

contrast we may consider blue-

bunch wheatgrass (Agropyron

spicatum). This species appar- ently has relatively low storage

concentrations, relatively long

basal internodes of 1% to 2

inches in length, and slow de-

velopment of abundant photo-

synthetic tissue (McIlvanie,

1942; and Branson, 1956). As

compared with crested wheat-

grass, it seems reasonable that

bluebunch wheatgrass would re- quire greater depletion of stored

reserves during initial growth

and require more time for its restoration.

After the early basal leafiness

947 993

12-17. This removed an aver-

was produced by crested wheat- grass, the clums arose rapidly

and uniformly. The culms ter-

minated growth by flowering

time.

In the consideration of carbo-

hydrate accumulation it is also

important to recognize environ- mental factors. Nitrogen fertili-

zation and other factors that

stimulate growth activity tend

to mobilize carbohydrates.

Graber (1931) reported that ni-

trogen fertilization may hasten

the decline of reserves under

close grazing or clipping. Bene- dict and Brown (1944) reported similar results regarding the ef- fects of nitrogen fertilization. Several workers have found the

rate of carbohydrate accumula-

tion to be inversely related to

the rate of growth. The infer- ences are that patterns of car- bohydrate depletion and accum- ulation may be quite variable from year to year and site to

site; although related to mor-

phological and phenological fea-

tures that characterize different species.

In brief, a plant’s demand for

carbohydrates in respiration,

growth, and reproduction must

276 D. N. HYDER AND F. A. SNEVA

stored reserves and current

photosynthetic production. Pro-

duction in excess of demand pro- motes active storage, and vice versa. Crested wheatgrass, with

relatively high storage levels,

short basal internodes, and early

abundant leafiness, developed

high photosynthetic production

rapidly while demands in stem elongation remained low. Some- what later, when reserves and

photosynthetic production were

high, the additional burdens in

growth and reproduction were

not especially serious. Within

this framework of carbohydrate

supply and demand there is op-

portunity to recognize species

and environmental differences

that are important in grazing

management.

The decrease in carbohydrate

concentrations at about flower-

ing time was intriguing because data on other species sometimes

indicate similar decreases. It

seemed logical to assume that

the energy expended in flower- ing was directly involved in the

requirement for utilization of

stored carbohydrates. However,

deheading did not affect carbo- hydrate trends, and it must be

concluded that flowering w a s

not the direct cause of the tem- porary decrease in food reserves.

The requirements in very fast

stem elongation after head

emergence were likely dominant

in producing the imbalance be-

tween carbohydrate production

and utilization.

The data presented on the

growth and carbohydrate trends

in crested wheatgrass support

the contentions of early range

readiness and high tolerance to grazing often attributed to this species.

Summary

Herbage yields and root-carbo- hydrate concentrations in crested

wheatgrass (Agropyron deser-

torum) were obtained in 1956 and 1957. In 1956 mature potted plants were used. Four plants were removed every 2 weeks to

obtain herbage yields, root

yields, and r o o t-carbohydrate

concentrations. In 1957 plants

were taken directly from the

field on 2 sites. On site I herb- age and roots were removed ev- ery 2 weeks to obtain herbage yields and root-carbohydrate concentrations. On site II, trends in herbage yields and root-car-

bohydrate concentrations were

taken at weekly intervals on un- treated and on deheaded plants.

Deheading was accomplished by

cutting the culms at the base of

the heads shortly after emer-

gence from the boot.

Growing-season trends in the

carbohydrate content of roots

(with stem bases) were charac-

terized by: (a) An early ac-

cumulation of carbohydrates to

a level near 27 percent by the

time of head emergence, (b) a

moderate decrease at or just be-

fore flowering that was not al-

tered by deheading, and (c) a

final recovery during or just fol- lowing seed formation to a level

approximately the same as that

attained by head emergence.

Within the framework of car-

bohydrate supply and demand

there is opportunity to recognize

species and environmental dif-

ferences that are important in

grazing management. Some phe-

nological and morphological

characteristics of crested wheat- grass were discussed as a basis

for the appreciation of species

differences in carbohydrate de-

pletion and accumulation. The

data presented on the growth

and carbohydrate trends in

crested wheatgrass support the

contentions of early range readi- ness and high tolerance to graz-

ing often attributed to this

species.

LITERATURE CITED

BENEDICT, H. M. AND G. B. BROWN. 1944. The growth and carbohy- drate responses of Agropyron smitthii and Bouteloua gracilis to changes in nitrogen supply. Plant Physiol. 19: 481-494.

BRANSON, F. A. 1953. Two new fac- tors affecting resistance of grasses to grazing. Jour. Range Mangt. 6: 165-171.

. 1956. Quantitative effects of clipping treatments on five range grasses. Jour. Range Mangt. 9: 86-88.

BROWNE, C. A. AND F. W. ZERBAN. 1941. Methods of sugar analysis. 3rd ed., New York: John Wiley 8~ Sons, Inc., p. 836.

COOK, C. WAYNE AND L. A. STODDART. 1953. Some growth responses of crested wheatgrass following herbage removal. Jour. Range Mangt. 6: 267-270.

GRABER, L. F. 1931. Food reserves in relation to other factors limiting the growth of grasses. Plant Physiol. 6: 43-72.

MCCARTY, E. C. AND RAYMOND PRICE. 1942. Growth and carbohydrate content of important mountain forage plants in central Utah as affected by clipping and grazing. U.S.D.A. Tech. Bul. 818.

MCILVANIE, S. K. 1942. Carbohydrate and nitrogen trends in bluebunch wheatgrass, Agropyron spicatum,

with special reference to grazing influences. Plant Physiol. 17: 540- 557.

NEILAND, B. M. AND J. T. CURTIS. 1956 Differential responses to clipping of six prairie grasses in Wisconsin. Ecology 37: 355-365.

SAMPSON, A. W. AND E. C. MCCARTY. 1930. The carbohydrate metabol- ism of Stdpa pulchra. Hilgardia 5

(4) : 61-100.

SULLIVAN, J. T. AND V. G. SPRAGUE. 1953. Reserve carbohydrates in orchard grass cut for hay. Plant Physiol. 28: 304-313.

Fertilization of Native Range in the

Northern Great Plains

ROBERT W. LODGE

AgriculturaZ Research Officer, Experimental Farm, Swift

Current, Saskatchewan, Canada

Research in the use of fertiliz- ers on native range forage in the Northern Great Plains is being conducted to find if this may be

a method of increasing the re-

turns per acre. Clarke, Tisdale,

and Skoglund (1947) reported

the long term effect of manur- ing. An area of Bouteloua-Stipa

vegetation, manured in 1928,

sampled from 1933 to 1938, in-

clusive, yielded an average of

850 pounds per acre against an average of 420 pounds per acre from adjacent unmanured range.

Work conducted at the Range

Field Station at Cottonwood,

South Dakota, by Westin, Bunt- ley, and Brage (1955) from 1952 to 1954 showed increased yield on native range after fertilizer

applications. Using rates of 20,

40, and 80 pounds of nitrogen per acre on pasture that had been grazed at heavy, moderate, and light intensities, they found the greatest response resulted from

the application of 80 pounds of

nitrogen per acre to the heavi-

ly grazed pasture. Increases in

percent protein were obtained

from the higher rates of nitro- gen. Rogler and Lorenz (1957)

conducted a fertilizer experi-

ment on an upland blue grama

(Bouteloua gracilis) -western

wheatgrass (Agropyron smithii)

site. Both heavily grazed and

moderately grazed range showed an increase in yield after appli- cations of 30 and 90 pounds of nitrogen per acre, with a greater return in pounds of hay pro- duced per pound of nitrogen at the 30-pound rate. Crude protein level was higher in the forage from the plots which received 90 pounds of nitrogen per acre than

from unfertilized plots.

These and other range ferti- lization experiments have shown that increased yields of forage may be obtained, and the ferti- lized forage will contain a great- er percentage of crude protein. They do not show that the in-

creased yields of forage and

crude protein are sufficient to

cover the costs of application.

Further certain inconsistencies

in the results suggest that the response of native range grasses

to commercial fertilizers needs

further investigation before

clear-cut recommendations can

be made as to their use as a range

management device.

This article reports a fertilizer

test on Stipu-Boutelouu range

near Swift Current, Saskatche- wan. The results of the experi- ments in themselves verify the findings of other work reported.

However, observations made

during and subsequent to the

test indicate that the measure- ment of certain reactions of the vegetation complex and of indi-

vidual species to fertilization

might be desirable.

Procedure

In 1951 a test was laid down to determine the effect on yield and on protein content of a single ap- plication of fertilizers and ma- nure to mixed-grass prairie. The test was located on the Alexan- der Coulee sheep range, Experi-

mental Farm, Swift Current,

Saskatchewan. The range was in

excellent condition prior to the

experiment, and the area used

was not grazed during 1951 and

1952. Needle-and-thread (Stipu

comutu) and blue grama are the

277

dominant grasses, other major

species being western wheat-

grass, Junegrass (Koeleriu cris-

tutu) and threadleaf sedge

(Curex filifoliu). The soil of the site is classified as Haverhill Loam, rolling phase of the chest- nut or brown soil zone, devel-

oped over undifferentiated gla-

cial till (boulder clay) . The cli-

mate is semi-arid, the long term

precipitation at Swift Current

being 14.94 inches, the average seasonal rainfall (May-June, in- clusive) is 6.98, and the precipi-

tation-evaporation index is 0.45.

Annual precipitation was above

average prior to and during the study, being in 1951, 19.06 inches and in 1952,16.05 inches, and the

seasonal rainfall was 5.17 and

8.58 inches in 1951 and in 1952.

The experimental layout was

a randomized split plot with six replicates. Plots were 3 x 14 feet. Treatments consisted of four fer-

tilizer treatments and one ma-

nure treatment each at two rates, and a check which was not fer-

tilized or manured. Fertilizers

used were ammonium phosphate

16-20-0, ammonium phosphate

11-48-0, ammonium nitrate 33.5

O-O, and barnyard manure. These

were hand broadcast in April,

1951, at rates of 16 and 32 pounds per acre of nitrogen for the arti- ficials, and 10 and 20 tons per acre of manure. The plots were mowed in late July, 1951, and early August, 1952, and the for- age from a central 18-inch wide

strip was collected from each

plot, air dried and weighed. The

samples from each treatment

were later bulked, ground, and analysed for protein content.

Resulfs

Forage yields obtained in the two years are given in Table 1. There was no increase in forage yield from the fertilizer applica- tions in 1951. In 1952 there was a significant increase in yield as a

result of the application of 32

278 ROBERT W. LODGE

Table 1. Yield in pounds of forage per acre (dry mafier) of mixed-grass prairie fertilized or manured at two rates.

-

1951 1952

16 1bs.N. 32 lbs. N. 16 lbs. N. 32 lbs. N. Treatment or 10 tons or 20 tons or 10 tons or 20 tons of manure of manure of manure of manure

Ammonium Phosphate 16-20-o 341 290 620

Ammonium Phosphate 11-48-o 464 453 493

Ammonium Nitrate 33.5-O-O 302 412 600

Barnyard Manure 371 385 623

No fertilizer treatment 325 337 482

SD. between treatments means: 1952 at 5% level 118 pounds. at 1% level 234 pounds.

468 505 666 767 442

plots which received 20 tons of manure.

Results of analyses for crude

protein content are given in

Table 2. From these analyses and the forage yields the gross crude protein per acre for the various treatments was computed (Table 3).

In 1951 there were increases in crude protein content as a result of the treatments, and as a result

a significant increase when 20

tons of manure was applied. Sig- nificant increases in gross pro- tein also resulted from the ap- plication of 32 pounds per acre of nitrogen as 33.5-O-O and ll-

48-O. The application of 16

pounds per acre of nitrogen as 11-48-O gave a very significant increase in gross protein.

In 1952 the percent crude pro- tein of the forage from the treat- ed plots was not apparently dif- ferent from that of the plots which received no fertilizer, ex- cept in that from the plots which received 10 tons of manure. This is reflected in the 1952 data in Table 3 in which this treatment

(10 tons of manure per acre) as well as the 32 pounds per acre of nitrogen application (as 33.5-O-O) ,

gave a signif icant increase.

Twenty tons of manure gave a

highly signif icant increase in

gross protein.

Discussion

Range fertilization in this

study is concerned with relative- ly low rates of application broad- cast in the spring. While drilling in of fertilizers is acknowledged

superior to broadcasting, the lat- ter is a more practicable method

for rough, rolling, or stony

ranges. Whether spring applica-

tion is advantageous over fall

application is not demonstrated. The differential effect of the sev- eral artificial fertilizers indicat- ed that the increase in percent protein may be in part due to the provision of phosphorus, but all treatments gave an increase in the quality of the forage in the

year of application. Yield in-

creases cannot be expected from the use of fertilizer at rates of the order of 16 pounds of nitro- gen. When 32 pounds of nitrogen were applied, the forage yield

increased in the second year

after application.

The use of manure in this ex- periment was intended to allow a comparison of it with those of Clarke et al. (1947). The first and second year advantages of the use of manure at rates of 10 and 20 tons per acre are appar- ent, both rates of application re-

sulting in increased gross pro-

tein per acre in the first year and increased forage yields in

the second year. On this some-

what more productive range

type than that studied by Clarke the value of manure is as appar- ent in the short term as were the

long-term advantages reported

by Clarke. The use of supplies

of manure from feedlots and

wintering lots on selected areas of native range can have immedi-

ate as well as long continued beneficial results.

Continuing studies in the use of fertilizers on the native grass

ranges of the Northern Great

Plains are warranted.

Kilcher (1958) reported on the response to fertilization of three species of cultivated grasses. His

work and the observations of

Rogler and Lorenz (1957) on

native prairie indicate the differ- ential response of species to addi- tions of nitrogen. Further studies

should be concerned not only

with the response in yield and in chemical composition of individ- ual species, but more important they should examine the result-

ant changes in botanical com-

position of the sward.

Many workers in forage crop

improvement programs have ex-

amined the effects on grass seed production resulting from appli- cations of fertilizers. It is pos- sible that in the complex of spe- cies that is a natural grassland, there might occur a differential response of the several dominant species. This certainly occurs in

sexual reproduction, and possi-

bly in vegetative reproduction.

Rogler and Lorenz (1957) imply

the latter. Moreover, the re-

sponse to fertilization of individ-

Table 2. Percent crude protein of the forage harvested from mixed-grass prairie fertilized or manured at two rates.

1951 1952

16 lbs. N. 32 lbs. N. 16 lbs. N. 32 lbs. N. Treatment or 10 tons or 20 tons or 10 tons or 20 tons of manure of manure of manure of manure Ammonium Phosphate 16-20-o 10.05 10.36 8.45 8.08 Ammonium Phosphate 11-48-O 10.10 9.53 8.00 8.32 Ammonium Nitrate 33.5-O-O 10.00 10.38 8.24 8.13

Barnyard Manure 11.01 12.10 9.22 8.83

FERTILIZATION OF NATIVE RANGE 279

ual species in an overgrazed Table 3. Gross protein in pounds per acre of fhe forage harvested from sward might well be expected to mixed-grass prairie fertilized or manured at two rates.

differ from the response of these _{1951 1952}

same species in an undergrazed

and therefore more stable stand. 16 lbs. N. 32 lbs. N. 161bs.N. 321bs.N.

There appears the possibility Treatment _{of manure of manure of manure of manure} or 10 tons or 20 tons or 10 tons or 20 tons

that range fertilization might

enable manipulation of the bo- Ammonium Phosphate 16-20-o 34 30 52 36

tanical composition of native Ammonium Phosphate 11-48-o _{Ammonium Nitrate 33.5-O-O} 47 ₃₀ 43 ₄₃ 39 ₅₀ 42 ₅₄

ranges. But such manipulation Barnyard Manure _{41 47 57 68}

to serve best the nutritional re- No fertilizer treatment 28 30 41 37

quirements of the grazing ani- __. -___

ma1 will first require additional S.D. between treatment means: 1951 at 5% level 13 at 1% level 17 pounds. pounds.

attention to the effects of the change in nutrient status on the

1952 at 5% level 15 pounds. at 1% level 20 pounds.

individual plant species and on

the vegetation complex.

The warrant for any range

management technique is in-

creased animal production. A

fertilization test to be realistic

must be concerned with in-

creased animal gains. The work of Smith and Lang (1958) who report on the use of nitrogenous fertilizers in achieving distribu- tion is worthy of attention. The value of fertilization of a range may be reflected in other meas- ures than increased forage quan-

tity or chemical quality. The

evaluation of fertilizer applica- tions must include any benefits of better distribution, increased

palatability, or increased feed-

ing value. Further grazing trials in which the increased produc- tivity is equated to the costs of fertilizer and its application are particularly necessary.

Conclusions

The evidence presented indi-

cates that the benefits of heavy applications of barnyard manure

are both immediate and last-

ing, whereas the application of

3.

commercial fertilizers may or

may not produce sufficient extra growth to warrant their use.

However, the observations

made during the course of this

experiment and the suggestions

indicated by literature reviews are such that certain confusing

issues are apparent. It is be-

lieved that further research

should consider the following

points:

That the study of the re- sponse of individual species

to a specific mineral or

combination of minerals

should be a feature of fer- tilization studies.

The response of individual species and changes in bo-

tanical composition of the

fertilized sward, should be interpreted only after con- sideration of the influence of different degrees of graz- use, past and future. That the effects of range fertilization on the growth

and development of the

livestock using the range

be considered. It is not the

increased yield of forage or crude protein that is the ultimate concern, but that

application of the knowl-

edge will mean bigger

calves and lambs and more wool and beef.

LITERATURE CITED

CLARKE, S. E., E. W. TISDALE AND N. A. SKOGLUND. 1947. The effects of climate and grazing practices on short-grass prairie vegetation. Can. Experimental Farm Service. Pub. 747. Tech. Bulletin 46: 51-52. KILCHER, M. R. 1958. Fertilizer

effects on hay production of three cultivated grasses in southern Sas- katchewan. Jour. Range Mangt. 11: 231-234.

ROGLER, G. A. AND R. J. LORENZ. 1957. Nitrogen fertilization of Northern Great Plains rangelands. J o u r . Range Mangt. 10: 156-160.

SMITH, D. R. AND R. L. LANG. 1958. The effect of nitrogenous ferti- lizers on c a t t 1 e distribution on mountain range. Jour. Range Mangt. 11: 248.

Effects of Presowing Vernalization on Survival

and Development of Several Grasses

NEIL C. FRISCHKNECHT

Range Conservationist, Intermountain Forest and Range Experiment Station, Forest Service, U. S. Department of Agriculture, Ogden, Utah

Improvement of plant estab-

lishment, especially on poor

sites, is an important problem in seeding depleted watersheds and ranges in the Intermountain West. One approach to this prob- lem lies in plant stimulation for

more rapid development to en-

hance the young plant’s chances for survival. The effectiveness of

presowing vernalization of seeds

in producing this kind of stimu- lation in some perennial grasses

was investigated in experiments

reported here. The procedure in- volves storing soaked seeds at near-freezing temper’a t ur e s f 0 r several weeks as a conditioning process for subsequent phasic de- velopment and flowering, as dis- tinguished from mere growth.

These studies are a follow-up of earlier work (Frischknecht,

1951) which suggested that in

certain grasses fall planting stim- ulated faster growth and devel- opment than spring planting, aside from the fact that seedlings

began growing earlier in the

spring. Mountain rye (Secale

montanum), intermediate wheat-

grass (Agropyron intermedium),

and a native strain of mountain

brome (Bromus carinatus) were

3. of 16 grasses studied which

merely stooled from spring

plantings, but they flowered and

produced seed the first year

from late fall plantings (Figure 1) even though seedlings did not

emerge until spring. Inasmuch

as this behavior parallels reac-

tions in so-called “winter” ce-

reals, it was believed possible to

“vernalize” seed of some peren-

nial range grasses and thereby

speed the development of spring

plantings. It appeared that this could be done underneath snow, especially inasmuch as soil ther- mograph records had shown that

winter temperatures near the

ground surface under snow re- mained close to 32” F., which were similar to temperatures used in vernalizing cereals.

According to the symposium

by Murneek and Whyte (1948)

vernalization of seeds was con-

sidered to have great practical value in Russia at one time for

hastening maturity in annual

plants. Winter cereals had re- ceived most attention. However these authors concluded that ver- nalization would have little prac- tical value in countries not ex- periencing extreme conditions of frost, drought, and floods espe- cially when superior genotypes were found. An exception would

be in the production of market

garden crops where a few days’ earliness would mean increased

financial returns. Martin (1934)

considered the process to have

little practical value in the

United States. McKinney (1940)

noted that most investigators

outside Russia attached little

importance to the economic gain from vernalization of plants.

Investigations of vernalization of both plants and seeds have continued on a great variety of species, mostly in other ‘coun- tries. In this connection Wellen- siek (1952) gave four illustrations of control of flowering: (1) plants that are insensitive to low tem-

peratures and day length; (2)

plants that require cold when they have reached a certain veg- etative size (plant vernalization); (3) plants that react to seed ver- nalization; and (4) plants sensi-

tive to plant vernalization but

also to short day, provided it is

followed by long days. Sechet

(1953) grouped a number of

plants into three categories with respect to their requirement of a

period of vernalization as fol-

lows: (1) determinant, in which a cold period is valuable but not

always indispensable for repro-

duction (winter cereals, tulips); (2) favorably responding to cold, in which flowering is precocious but occurs without a cold pe-

riod (mustards, lupine, peas,

etc.); (3) no favorable response to vernalization (onion, flax, bean, etc.).

FIGURE 1. Mountain rye from spring planting (left) merely stooled during first growing

season, whereas fall planting (right) stimulated flowering and seed production, although

seedlings did not emerge until spring.

EFFECTS OF VERNALIZATION OF GRASSES 281

Some investigations, such as

the series by Gott, Gregory, and Purvis (1955), have been aimed

at discovering the fundamental

biological processes involved.

Most of the recent investigations in the United States have cent- ered around substances affecting

plant development and flower-

ing and control of the process by

their application. Associated as-

pects of photoperiodism have re-

ceived more attention than tem-

perature - the main factor in

vernalization. Little work has

been done on seed vernalization in perennial grasses.

*

Preliminary Attempt at Vernalization

The first of this series of studies, by the author, made in

central Utah, involved soaking

seeds of eight perennial grasses for 20 hours at room tempera- tures and then burying them in

a snowbank at the ground sur-

face for 50 days before planting. The grasses were mountain rye,

intermediate wheatgrass (regu-

lar and Amur strains), pubescent

wheatgrass (A. trichop horum),

tall wheatgrass (A. elongatum),

fairway wheatgrass (A. crista-

turn), crested wheatgrass (A. de- sertorum), and Russian wildrye (Elymus junceus).

Duplicate samples of seeds

thus treated and similar untreat- ed samples were planted at com- parable rates in early April 1952, at each of three locations in cen- tral Utah: (1) Benmore, big sage-

brush (Artemisia tridentata)

type, average annual precipita- tion 12.8 inches; (2) Tintic Valley,

big sagebrush type, estimated

average annual precipitation 10

inches; and (3) Gunnison, shad- scale (Atriplex confertifolia)

type, estimated average annual precipitation 8 inches. Seed sam- ples were planted in separate 5- foot rows, spaced 20 inches apart. The soil was moist on all sites

at the time of planting; this

helped to prevent the moist

snowbank-treated seeds from

FIGURE 2. Two years after spring planting at Gunnison good stands were present only

from snowbank-treated seeds of Russian wildrye (right foreground), crested wheatgrass

(third row left), and fairway wheatgrass (seventh row left). The other half of each row

had been planted with untreated seeds.

drying. A light rain fell at Gun- nison on the day of planting, and moisture fell at the other two sites on the third day after planting.

Seed germination tests imme-

diately following removal from

the snowbank showed that vi-

ability was not impaired, except in mountain rye. Many seeds of mountain rye were decomposing after 50 days in the snowbank;

similar damage was reported

previously for dehulled seeds of this grass, tall oatgrass, and or-

chardgrass overwintering in

cloth bags in the ground (Frisch- knecht, 1951). Apparently, if seed

is dehulled, planting of these

grasses should be done when

there is reasonable chance for

prompt germination.

Except for mountain rye,

treated seeds produced good to

excellent seedling stands on all

plots at all three sites. None of

the grasses produced seed the

first year, but plants from all

snowbank-treated seeds emerged

a few days earlier than plants from untreated seeds, and they grew taller the first year. This

was particularly striking in the

two strains of intermediate wheatgrass: their seedlings from

treated seeds tended toward

culm elongation instead of stool- ing like seedlings from untreated

seeds. A similar tendency was

observed on the few plants of

mountain rye that were present on each site from snowbank-

treated seed. Such development would suggest that vernalization had occurred in these two spe- cies at least.

The generally g o o d initial stands were maintained at Ben- more. Rabbits greatly damaged the plantings at Tintic Valley by the end of the first season. The

following spring only the two

rows of Russian wildrye and one

row of pubescent wheatgrass

from snowbank-treated seed,

plus one row of Russian wildrye

from untreated seed showed

good stands. High mortality oc- curred at the shadscale site, and by 1954 the only remaining good stands were from snowbank- treated seeds of Russian wildrye, crested wheatgrass, and fairway wheatgrass (Figure 2).

Snowbank versus Other Presowing Treatments

Other presowing treatments were compared with snowbank

treatment in a subsequent test

involving four selected species.

Presowing treatments of seed in- cluded: (1) 20 hours’ soaking at

room temperature followed by

snowbank storage for 48 days;

(2) 20 hours’ soaking followed by 48 days’ storage in a locker at

extremely cold temperatures

(near 0” F.); (3) 36 hours’ soaking followed by 3 days’ locker stor- age; and (4) 36 hours’ soaking fol- lowed by no cold treatment. Spe-

cies used were intermediate