Published July, 1962
Factors Affecting Tillering in Alfalfa'
Everett R. Cowett and Milton A. Sprague'
SYNOPSIS. Tillering of established plfalfa plants was
increased by anti-auxin application and a short photo-
period, whereas tillering of seedlings was increased by
:long photoperiods. Soil moisture, light intensity, tem-
perature, mineral nutrition, and cutting treatment altered
bud and stem development concurrently as they affected
vigor and over-all growth.
HE Occurrence of basal shoots on the crowns of alfalfa
T (Medicago Jutiua) plants is a natural phenomenon, one
which is largely responsible for vegetative recovery of the
species. Furthermore, in an area where individual plants
are sparse, rapid tillering commonly occurs, partially filling
in the open areas between plants. Also, an increase in the
number of stems per plant is often accompanied by in-
creased yields, IeafJstem ratios, and better-quality forage.
This investigation examines the extent to which tiller
formation is altered by environmental and physiological
factors. Particular emphasis is given to the effects of soil
moisture, light intensity, air temperature, nutrient level,
growth regulators, photoperiod, cutting height, maturity at
harvest, and stand density on bud and tiller development.
LITERATURE REVIEW
The control mechanisms which give rise to buds to form
the characteristic crown are not clear. Very commonly, 3 or
4 branches, in addition to the primary axis, form the
crown of the alfalfa seedling ( 4 ) , whereas older plants
may have 5 to 20 or more stems in their crown. Jones ( 5 )
found alfalfa varieties to vary in crown development, along
with vigor of the plant and the environment. However,
little data is available specific to those factors affecting
tiller and crown development of alfalfa.
Of the conditions observed by Willard (11) dry weather
was the only one that retarded the development of crown
buds. Day length had no effect on number of stems or
buds of alfalfa seedlings, but. long photoperiods were con-
ducive to higher yields (7). Leopold (6) found additions
of auxin to barley p!ants resulted in changes in tillering
frequency. Similarly, Asen and Hamner ( 1 ) increased the
number of basal shoots of roses by applications of anti-
auxins. Thompson ( l o ) , however, did not alter the number
of stems on alfalfa plants with growth regulators but did
alter yield.
Yield of alfalfa and the development of crown buds and
shoots were found to be associated with total available car-
bohydrates in the roots at the time of the last cutting (2,
3 ) . Nelson (8) found, however, that crown bud, and tiller
formation accompanied frequent cutting and root reserve
depletion.
EXPERIMENTAL PROCEDURE
Except for seedling studies, vegetative cuttings obtained from a
single plant of Atlantic alfalfa were used. Cuttings were rooted
for about 14 days in sand initially saturated with water. After
rooting, cuttings were placed in 2- by 2-inch paper bands in green-
house flats containing a mixture of equal parts by volume of sand,
peat, and a Nixon loam. A similar media also was used in the
experimental pots. Immediately after planting, the soil was inocu-
lated with proper strains of nodule bacteria, sprinkling inoculum
over the surface and washing it in with water. Established plants
294
at heights of 4 to 6 inches were selected for uniformity and used
for experimental work. Seedlings of Atlantic alfalfa were obtained
from inoculated lots of breeder's seed planted in a Nixon loam.
Soil moisture and light intensitj-At 4 weeks of age, seedling
alfalfa plants growing in a Nixon loam soil were subjected to
approximately 10, 3 , and 1/2 atmospheres of moisture tensi,m (low,
medium, and high), and 4 light intensities (25, 50, 75, and 100%
of full sunlight). Three replications of each treatment were pro-
vided. Moisture stresses were maintained by weighing each pot
daily and adding sufficient water to obtain the weight desired.
Light intensities were provided by canopies considng of
wooden frames over which were drawn numbers of layers of white
cheese cloth chosen to permit desired percentages of light to pass
through.
Treatments were maintained for 4 weeks, after which plants
were harvested and their heights, weights, and numbers cd stems,
buds, and leaves per plant were determined. Root weights were
obtained from plants in the full sunlight treatment grown at 3
moisture levels. Pots were rearranged at this time so thai: 3 pots
of each watering treatment remained at the same level of moisture
and 3 others were transferred to each of the other moisture
regimes, all under full sunlight. The pots were so arranged that
each of the light intensities prior to the first harvest now repre-
sented one replication of the new arrangement. Following it second
4-week period of growth, plants were again harvested and similar
data recorded.
Temperature-Ten clonal cuttings were selected for ea<:h of 2
temperature treatments and transplanted into individual 5-inch
pots. Cuttings were grown for 2 weeks in the greenhouse before
subjecting them to day-night temperatures of 75 to 60 and 90 to
75" F. in growth chambers. Lighting was supplied by cool-white
fluorescent tubes providing an intensity at plant height of 1,000
foot-candles for 16 hours of each day. Plants were harvestcd twice
at 4-week intervals during their growth in the chambers.
Mineral nutrition-Twenty-two alfalfa seedlings were established
in each of twenty-four 2-gallon pots, filled to 1 inch from the top
with washed quartz sand. Nutrient solutions for use in the sand
culture were formulated to provide 8 treatment combinations of
high and low levels of nitrogen, phosphorus, and potassium. The
elements were maintained at 140, 62, and 156 parts per million
to represent high levels and 21. 0.92, and 2 3 parts per million to
represent the low levels of N, P, and K, respectively. Minor ele-
ments were provided in abundance uniformly for all tre-tments.
Each treatment appeared three times in a randomized block design.
A substantial quantity of the appropriate nutrient formulation was
applied to each pot daily from the day of planting until hazvest.
Plant-gvowth vegulators-Triiodobenzoic acid (TIBA) at 5, SO
and 500 micrograms per plant, naphthleneacetic acid ("PA) at
500 micrograms per plant, and a control treatment receiving only
lanolin paste were studied for their effects on tillering in the
greenhouse. Three replications of 1 plant each were provided using
clonal cuttings. When plants were 2 5 cm. tall, growth substances
were mixed in lanolin paste and applied by encircling the main
stem of each plant with a given material just above the crown.
A Luer syringe permitted a uniform application of 0.05 ml. of
lanolin containing the desired quantity of growth substance per
plant. For ease of application the paste was warmed gently until
it flowed easily. After 6 weeks, plants were cut just above the
area at which the growth substances were applied and all active
growing points removed. Numbers of stems, buds, and dry weights
were determined, and the plants permitted to recover for a 5-week
period, at the end of which time observations were repeated.
Photoperiod-Four-week-old alfalfa seedings and vegetative cut-
tings 6 inches tall were compared under 10- and l d h o u r photo-
periods in the greenhouse. Nine 9-inch pots, of 20 seedlings each,
were used for each treatment. In addition, 9 vegetative cuttings
'Paper of the Journal Series, Department of Farm Crops,
Rutgers-the State University, New Brunswick. N. J. Rcceived
Sept. 14, 1961.
Formerly Research Assistant, now Assistant Agronomisi, Uni-
versity of New Hampshire, Durham, and Professor of Farm Crops,
Rutgers, respectively.
 COWETT & SPRAGUE: FACTORS AFFECTING TILLERING I N ALFALFA 295

in individual pots were provided for comparison. All plants were Table &Recovery growth after cutting of alfalfa cuttings
maintained in desired environments until plants under long days treated with triiodobenzoic acid (TIBA) and naphthalene-
had flowered, during which time three successive harvests were acetic acid NAA).
taken from plants under each treatment.
Growth regulator Characteristic of growth
Cutting height-Nine vegetative cuttings were selected for test- treatment
Gtems Stems Plant Drywt. D r y wt.
ing each of 3 heights of cutting (1, 2 , and 3 inches) whereby 0, andbuds per height, tops, roots,
1, and 2 branch bases were left remaining at the base of stems. per plant plant cm. g. g.
Cuttings were made at full bloom and plants permitted to recover 5pg. TIBA I. 0 4.3 26.0 n. 62 0. 56
for 5 weeks in the greenhouse before a second harvest. 50 pg. TIBA 6. I 3.3 36.5 1.00 0.74
500pg. TIBA 8. 3 5.3 29. I n. 72 0.71
Stage of maturity-Nine vegetative cuttings were selected for 500pg. NAA 2.3 1.3 30.3 0. 56 0.63
clippings at early bud, early bloom, and past bloom. Plants were Lanolin (control) 3. I 2.3 45.7 1.24 0.16
clipped to their bare crowns at the desired stages of growth and LSD (0.05) 3.4 2.6 NS NS NS
permitted to recover until early bloom.
Stand density-Early in May, 1960, eighteen 4- by 4-foot plots Table 5-Influence of photoperiod on the development of alfalfa
were established on a uniform area of a Nixon loam soil. To this seedlings.
area, which tested medium in general fertility, 600 pounds per
acre of 5-10-10 fertilizer were applied and the soil limed to a 3 n g t h of Characteristic of growth
p H of 6.2. After fitting the soil, 1,170 vegetative cuttings, 4 inches
tall, were selected from a single clone propagated in the green-
photoperiod
Sterns per
plant
Plant heieht.
cm.
I , Drv
~~
---~
---
, weirht
-,
per pot, g.
house and transplanted to the field in a systematic manner. Six 16 hours 6.P 19 '
38. ?*
2 0
. 1. 30*
1.30*
10 hours
plots of each of 3 stand densities, 1, 4, and 8 plants per square 3.5
-.I 20.4
* Y . l 0.61
foot, were established. At a time when growth had reached about * slgnificmt at 5% level.
*slgnificmt

10 inches, the main stems were removed and discarded. Six weeks
later each plant was harvested individually and numbers of stems Light intensity-The magnitude of differences in the
and buds and dry weights determined. development of alfalfa seedlings under different light in-
tensities was small (Table 2 ) . High light intensities pro-
RESULTS duced greater numbers of stems per plant and higher yields
Soil moisture-Soil moisture expressed itself as a factor than low light, but differences were not significant above
controlling over-all growth as all characteristics varied and below 50% of full sunlight. Plant height was favored
similarly (Table 1). Plants with the greatest yields of either by low light intensity whereas the number of buds was not
tops or roots, and those with the greatest height possessed affected.
the largest number of stems and buds per plant. Soil mois- Temperature-No significant differences in the number
ture during a previous growth period had no significant of stems per plant or in heights of vegetative cuttings were
effect on plant growth at the later harvest. The number of apparent under 2 temperature regimes at 2 successive har-
stems was greater where soil moisture was high during the vests. However, lower day-night temperatures produced
recovery period regardless of the size of the crowns or more dry matter at both harvests and a larger number of
root system at the previous harvest. stems and buds combined at the second harvest. Further,
plants maintained at high temperatures did not recover
Table 1-Growth of alfalfa seedlings as influenced by soil after the second harvest, whereas those at the lower tem-
moisture. perature recovered well.
Moisture Characteristic of growth
Mineral nutrition-Greatest tiller development and yield
Sterns Buds Plant Rootwt. Top w t of both tops and roots occurred under high levels of nitro-
per
plant
per
plant
height,
cm.
per pot,
g.
per pot,
g.
gen, phosphorus, and potassium (HHH). This was fol-
LOW 2.58 1.22 18.0 2.14 2.19 lowed by treatment HHL, and treatment LLL, respectively
Medium 3. 33 1.54 36.4 4.14 5.13 (Table 3 ) . Significant differences between some of the
High 3. 56 1.18 42.5 4.41 6. I1
LSD (0.05) 0.24 0.52 4.5 1. I9
other treatments were less marked, but indicated possible
additive effects and the benefits of balanced nutrition.
Particular attention should be given to treatment LLL. This
Table 2-Growth and development of alfalfa seedlings as influ- treatment, low in all three elements, produced a greater
enced by light intensity. number of stems, abundant buds, greater plant height, and
Percent of Characteristic of growth
higher yields than other treatments less well balanced
full sunlight Stems per Buds per Plant height, D r y wt. except for treatment HHL.
plant Plant crn. per pot, g. Plant-growth regulators-Three rates of anti-auxin (tri-
100 3.31 1.34 21.6 5.42
75 3. 32 1.61 34.3 5.11 iodobenzoic acid), one rate of auxin (naphthaleneacetic
50
25
2. 81
2.83
1.60
1.46
32.8
38.4
4.38
3. I9
acid), and lanolin alone affected the first growth of alfalfa
ISD (0.05) 0.41 . NS 2.8 0.17 cuttings similarly. Branching of the main stem and absence
of basal shoots of the first growth were the same irrespec-
tive of treatment applied. Re-growth, however, revealed
Table 3-Eff ect of nitrogen, phosphorus, and potassium levels
o n growth of alfalfa seedlings in a sand culture. striking differences. Anti-auxin increased tillering and bud
development at all rates while the auxin inhibited the
N PK Characteristic of growth development of buds and shoots (Table 4).
levels Sterns Buds Top wt. Crownwt. Rootwt. Plant
per per per pot, per pot. per pot. height.
Photoperiod-Alfalfa seedlings subjected to 16 hours of
+
plant plant g. g. g. crn. light produced more dry weight, were taller, and possessed
HHH
HHL
I. 2
5. 9
2.5
1.8
23.25
16.18
3.17
1.56
29.07
14.16
32.4
26.4
more stems, active buds, and leaves per plant than plants
HLR 1.6 0.4 1.14 0.22 1.32 5. a grown under 10 hours daylength (Table 5 ) . Vegetative
L" 3. 0 1.1 1.58 0.22 2.46 a. n
LLH 1.3 0.4 0.87 0.15 1.54 5. I cuttings and established plants, by contrast, were affected
LHL
HLL
2.8
1.4
1.4
0.2
1.65
0.85
0.26
0.23
2.40
1.49
8.8
4.7
by day length quite differently than seedlings (Table 6).
LLL 4.6 1.1 4.46 0.49 5.33 14.2 During the first growth of vegetative cuttings only dry
LSD (0.05) 1.1 0.6 3.18 0.39 1.32 3.0 weights were significantly affected. At later harvests plants
296 AGRONOMY JOURNAL

Table &Effect of photoperiod on vegetative cuttings and on Table ?'-Regrowth of alfalfa as influenced by height of cut.
regrowth following harvest of alfalfa.
Cutting Characteristic of growth
h g t h Of characteristic of growth height Stems Stems and Stems Stemsand Flowers Plant Dry
on buds on on buds on per height, weight
i1ho)ope*id Stems per Buds per Plant height, Dry weight entire entire true true plant cm. per plant,
plant plant cm. per plant, g. plant plant crown crown g.
1st harvest: 16 hr. 2.1 0 63.8 0.84* 1-inch 4.2 7.4 4.2 7.4 5.7 70.0 2.71
10 hr. 2. 3 0 57.2 0.62 2-inches 5.3 9. 6 3. 7 6.8 10.8 74.6 3.60
Zndharvest: 16 hr. 2.9 5.1 58.8* 1.95* 3-inches 7.8 15.4 3. 9 7.8 23. 9 73.1 4. 81
10 hr. 3. 6 8. O1 41.0 1. 02 LSD(O.05) 1 . 5 4.3 NS NS 9.4 NS 1.33
3rd harvest: 16 hr. 4.3 1. 8 77.7* 2.971
10 hr. 5.2' 5.6" 47.1 2. 36
* Significant at 5 % level. Table 8-Effect of stage of maturity at harvest on the numbers
of buds, stems, and dry weight per plant at the following
harvest.
grown under a shorter day length developed a greater
number of stems and active buds per plant, but were shorter Stage of maturity
at 1 s t harvest
Characteristic of growth
Stems per Stems and buds D r y weight
and produced lower yields than plants under the long day plant per plant per plant,
treatment. Early bud 2.6 5.1 2.27
Early hloorn 3. 0 5. 8 2. 25
Catting height-Alfalfa cut high produced more stems Past bloom 3. 7 8.4 3.18
and buds on the entire plant but maintained the same num- I S D (0.05) 1.0 2.3 0.48
ber of stems and buds on the old crown (Table 7 ) . Greater
dry weight was produced and maturity was hastened when Table 9-Influence of three stand densities on growth of alfalfa
plants were cut high compared with cutting low. in the field.
Stage of maturity-Alfalfa cut at an advanced stage of
Plants per Characteristic of grewth
maturity produced more dry weight and more stems and square foot
Stems Buds Dry weight Total yield
buds per plant at a subsequent harvest than when cut early per plant per plant per plant,g. (T. / A , )
(Table 8 ) . No significant differences were obtained when 1 7. 1 2.4 22.0 1. 05
4 5.1 2. 3 10.6 1.95
comparing cuts at bud and early bloom stages. 8 4.0 1.8 7. 0 2.42
Stand density-As the stand density increased from 1 to LSD (0.05) 0. 7 NS 4.6 0.41
8 plants per square foot, the number of stems and dry
weight per plant decreased (Table 9 ) . As stand density
increased, however, the yield per acre increased. the three major elements produced higher yields and a
larger number of stems than treatments receiving higher
but unbalanced nutrient levels. It is possible that a bal-
DISCUSSION anced supply of nutrients may have an effect on e.tKciency
Soil moisture, light intensity, temperature, mineral nutri- of photosynthesis. Low phosphorus had more influ(-nce on
tion, and cutting treatments all affected the number of stems stem and bud development than other elements teskd. The
per plant only as they affected vigor and over-all growth fewest buds and stems were produced at low phosphorus
of the individual plant. Treatments which produced the levels, while the response to potassium was not as great.
greatest top and root growth, plant height, and number of These phenomena need to be studied more critically.
leaves also provided the largest number of stems per plant. The development of stems on alfalfa independent of
Carbohydrate production and accumulation as a ready growth occurred in only two instances. Short photoperiod
source of energy appears to be important for crown bud and anti-auxin applications increased the number of buds
initiation and development. This view is supported by the and stems per plant of aftermath with a concurrent: reduc-
fact that those treatments applied which would be expected tion in yield. Short photoperiods did not, however, increase
to either increase or decrease food reserve contents in roots the number of stems on alfalfa seedlings. Anti-auxin did
and crowns had a pronounced effect on crown bud initia- not affect the first growth of vegetative cuttings. Generally,
tion and tillering. Plants grown under high temperatures growth regulators and photoperiod treatments affect apical
had a number of stems at their first harvest comparable to dominance and subsequently the production of buds, stems,
plants grown under low temperatures, but their yields and and perhaps the number of branches on main stem::. Prob-
number of stems and buds were greatly reduced at later ably crown buds and stems of the second crop responded
harvests. Food reserve exhaustion, as described by Sullivan to level of storage of food reserves in the first crop and
and Sprague ( 9 ) , would be expected to have a bearing on were influenced by plant homones and photoperiod.
this response. Stage of maturity when harvested had an influence on
Alfalfa seedlings grown under moisture stress made very the number of stems per plant. Under the conditions of
little growth and had very small crowns and root systems this experiment, the effects were small and suggested that
at harvest. Yet, when these drought-conditioned plants were maturity was not of major importance. Repeated clippings
cut and subjected to higher moisture levels, comparable during successive crops might be expected to exert accu-
yields and stem numbers were obtained with plants having mulative effects. Cutting stems high at harvest increased
larger crowns and root systems but not similarly treated. both yield and number of stems and buds per plant at the
Apparently, an accumulation of carbohydrates in the roots following harvest. Plants cut high had the appearance of
and crowns under the low-moisture conditions (12) was having several crowns, whereas those cut short had a more
such that plants following this treatment produced more compact crown. Tiller development above the true crown
stems and buds per unit of crown than plants not subjected occurred only at the bases of branches left intact at cutting.
to moisture stress. Stand density in field plots exerted the most pron,ounced
Nutrient balance was revealed to be more important for affect on the number of stems per plant of any treatment
growth and tiller development of alfalfa seedlings than applied. Since changes in stand density also markedly
nutrient level. A treatment which received low levels of changes the moisture, light, and temperatures in the micro-
 COLVILLE: EFFECT OF RATE AND METHOD OF PLANTING ON YIELD OF IRRIGATED CORN 297

environment, the influence of stand on bud development is

not likely to be the control mechanism, but is accumulative
with respect to all factors involved. Further, it is likely
that some influence of stand on growth has not yet been
observed.

SUMMARY
Experiments were conducted in the greenhouse, growth
chambers, and field to evaluate factors affecting tiller pro-
duction in alfalfa. Soil moisture, light intensity, tempera-
ture, mineral nutrition, and cutting treatment altered bud
and stem development concurrently as they affected vigor
and overall growth. Tillering of established plants was in-
creased by anti-auxin applications and a short photoperiod,
but yields were decreased. Tillering and yield of alfalfa
seedlings was increased by long photoperiods.
Cutting alfalfa high or at advanced stages of maturity
increased yield and total number of stems per plant. Cut-
ting high did not, however, increase the number of stems
near the soil surface.
Stand density had the most pronounced effect on the
number of stems on alfalfa plants of all the factors studied.
Lower stand densities resulted in less yield per unit area,
but a greater number of stems per plant. Plant vigor, as
influenced by carbohydrate accumulations and energy rela-
tionships in roots and crowns, is likely to be the most
critical factor governing stem production in alfalfa.