9 Alfalfa

D. K. Barnes
USDA-SEA and University of Minnesota
St. Paul

Alfalfa (Medicago sativa L.) is an important forage species for hay and
pasture. Although it originated in southwestern Asia, it is well adapted to a
wide range of climates and soils in the United States, where about 11 million
ha are grown annually. Alfalfa has been referred to as "Queen of the For-
ages" because of its high yields and feeding value. Between 1900 and 1975
more than 160 cultivars were developed for production in North America
(Barnes et al., 1977). Most of the newer cultivars were selected for improved
adaptation and multiple pest resistance.

I. PARENTAL MATERIAL

The genus Medicago is widely distributed and comprises an array of di-

verse species that are either annual or perennial. The most recent taxonomic
studies of the perennial species concluded that M. sativa is polymorphic.
Lesins and Gillies (1972) defined the complex as M. sativa-falcata-gluti-
nosa, and Gunn et al. (1978) designated it as the M. sativa sensu lato com-
plex. Crosses can be made among subspecies in the M. sativa complexes.
Crosses can be made with the cultivated tetraploids and wild diploids with-
out special preparation of the parents (Bingham, 1968).

II. PLANT CULTURE

A. Field

Successful alfalfa seed production under field conditions requires

proper insect control, adequate pollination, and the adjustment of cultural
and management practices to local conditions (Pedersen et aI., 1972). High

Copyright (c:i 1980 American Society of Agronomy-Crop Science Society of America, 677
S. Segoe Road, Madison, WI 53711. Hybridization of Crop Plants.

177
178 BARNES

seed yields require well-drained soils which are low in alkali and soluble
salts, and a rooting depth of 1.2 m or more. Gravelly and shallow soils
should be avoided. Irrigation practices should prevent severe plant stress
and promote slow, continuous flowering through the entire production
period without excessive stimulation of vegetative growth. Areas with low
relative humidities and moderate to high temperatures are preferred for
alfalfa seed production because of a low incidence of leaf diseases, long
periods of pollinator activity, and favorable harvest conditions. Applica-
tions of fertilizers to established stands in the United States and Canada,
and foliar applications of major and minor elements to seed fields in Cali-
fornia, usually have failed to increase seed yields.
A critical aspect of seed production is crop management so that the
flowering period coincides with the period of least competition from other
pollen sources, the greatest activity of pollinators, and the most favorable
weather conditions (Pedersen et aI., 1972). In some areas, these circum-
stances require removing the first and possibly the second growth for forage
and then producing seed on the subsequent regrowth.
Rincker (1976) reported that seed increases during the year of establish-
ment can be maximized by starting seedlings in the greenhouse and trans-
planting them to the field at a rate of 9,000 to 10,000 plants/ha. He found
that transplants yielded more than twice as much seed as direct-seeded
plants during the year of establishment; however, seed yields were similar
during subsequent production years.

B. Growth Chamber and Greenhouse

Successful alfalfa seed production in the growth chamber and green-

house requires photoperiod and temperature adjustment plus insect and
mite control. Most alfalfa cultivars are long-day plants (Bula and Massen-
gale, 1972), but cultivars differ in their quantitative response to photo-
period. Flowering is usually sparse at photoperiods of 12 hours or less, but
at 14 to 16-hour photoperiods most plants will flower. NittIer and Kenney
(1964) reported that the most profuse flowering was under continuous light.
They also reported that flowers increased when light level was increased
from 12 to 24 p. E sec-I m- 2 under photoperiods long enough to promote
flowering.
Elgin and McMurtrey (1977) compared alfalfa seed production with
four types of supplemental lighting (high-pressure sodium, metal halide,
mercury vapor, and incandescent) at an 18-hour photoperiod. The amount
of flowering and subsequent seed production was directly related to the
quantity of light supplied. The most effective light source was the high-
pressure sodium lamp.
Temperature is another factor that can influence flowering in alfalfa
(Massengale et aI., 1971). Greenhouse and growth chamber conditions most
commonly used for alfalfa seed production include temperatures of 24 to 30
C and at least 16 hours of light.
ALFALFA 179

III. FLORAL CHARACTERISTICS

Essentially all annual species are cleistogamous and are exclusively self-
pollinated. Generally, the perennial species require tripping, and will set
seed from either self or cross-pollination.
The flowers of perennial species are morphologically complex and have
a unique tripping mechanism. The alfalfa flower is a classic example of an
evolutionary adaptation for cross-pollination by insects (McGregor, 1976).
A detailed description of floral development and floral morphology
was prepared by Barnes et al. (1972). Alfalfa flowers are borne on simple
racemes, with about 10 flowers per raceme. At the base of each flower the
receptacle is enclosed by the calyx tube consisting of five undiverged sepals
terminated by five lobes. The receptacle serves as the base for the corolla,
pistil, stamens, and nectary. Alfalfa has a papilionaceous corolla that
consists of five petals: a large standard, two lateral wing petals, and two
fused keel petals (Fig. la). The 10 stamens form a tube in which 9 filaments
are fused. The 10th stamen is nearest the standard and is free (Fig. 1b). Fila-
ments alternate long and short so that, during development, anthers fit
tightly around the stigma in a double ring (Fig. 2a). The pistil consists of a
single carpel that develops a superior ovary, a smooth awl-shaped hollow
style, and a well-defined stigma (Fig. 2c). The ovary contains an average of
10 to 12 ovules. The nectary is located in the base of the receptacle (Fig. 1c)
and covers the area from the base of the pistil back to the base of the keel
petals (Teuber, Albertsen, and Barnes, 1977). The nectary is from 2 to 8
cells thick and contains 7 to 25 stomata from which nectar is exuded. A
nectar reservoir is located above the nectary.
Before pollination can be successful in alfalfa, the sexual column
(ovary and staminal column) must be tripped (released). This is usually
done by bees foraging for nectar and pollen. In some instances, the flower
can be tripped by environmental factors such as hot, dry winds, and rain.
Larkin and Graumann (1954) indicated that the two major forces involved
in the tripping mechanism are pressure exerted by the sexual column from
cells under tension at the juncture of the staminal tube and the keel, and the
restraining mechanism of the keel petals that cohere due to interlocking pro-
jections of cutinized tissue in the appressed petal surfaces. Tripping takes
place whenever the restraint of the appressed keel petal is reduced or be-
comes less than the pressure of the staminal column. Plants differ in ease of
tripping.
Four bud stages are recognized in alfalfa: straight bud, pointed bud,
hooded bud, and erect standard (Coffman, 1922). Anther dehiscence usual-
ly occurs in the pointed bud stage (Fig. 2b). A cuticular membrane forms a
continuous film over the stigma, thereby preventing pollination before trip-
ping. When tripping occurs, the stigmatic membrane usually ruptures as the
stigma strikes either the bee or the standard petal. Alfalfa pollen is sticky
and readily adheres to pollinating insects. If the stigma fails to come into
180 BARNES

contact with foreign pollen, its own pollen germinates and produces pollen
tubes. Self-pollinated alfalfa has fewer flowers that form seed pods and
fewer seed per pod than cross-pollinated alfalfa.
The alfalfa flower gives sufficient protection to keep the pollen viable
and the stigma receptive from the late bud stages until the flower begins to
wilt. The time flowers remain open varies among genotypes and environ-
ments from about 5 to 16 days. Alfalfa pollen has been successfully col-
lected and stored from 2 to 6 months in sealed vials (Hanson, 1961; Lehman
and Puri, 1967). Pollen storage has rarely been used because alfalfa is
perennial, has an indeterminate flowering habit, and can be easily propa-
gated vegetatively. With a combination of field and greenhouse facilities,
flowering plants can be produced throughout the year.

Fig. I-A, Alfalfa floret; B, cross section of the upper calyx area; C, cross section of the base
of the receptacle area. In A, B, and C: c, calyx; fs, free stamen; k, keel petal; nr, nectar
reservoir; ns, nectary and nectary stomata; 0, ovary; ov, ovule; s, standard petal; sc, stami-
nal column; and w, wing petal (courtesy of L. R. Teuber and M. C. Albertsen).
ALFALFA 181

IV. ARTIFICIAL HYBRIDIZATION AND SELF-POLLINATION

A. Equipment

The equipment needed for pollination is similar to that described for

clover (Chapter 16).

B. Preparation of the Female

Successful self and cross-pollinations can be made with any flowers

that are not wilted and that have fertile ovules and pollen. Cross-pollination
by hand without emasculation has often been used to produce seed for plant
breeding studies. For critical studies in which self-pollinated seed cannot be
tolerated, either the female parent can be emasculated before pollination or
a male-sterile plant can be used as the female parent. Anthers and pollen
can be removed from a flower during emasculation by either suction or
alcohol (Tysdal and Gar!, 1940). With both methods the standard petal is
clipped and the flower is gently tripped. With vacuum emasculation, a glass
tube, drawn to a I-mm tip and attached to a vacuum source, is used to re-

Fig. 2-Alfalfa reproductive development. A, alternating long and short stamens at straight
bud stage of development; B, dehisced pollen surrounding stigma at pointed bud stage of
development; C, mature ovary, style, and stigma.
182 BARNES

move the anthers. A low-powered magnifier can be used to determine

thoroughness of pollen removal. A skilled operator can expect normal seed
set (three to five seeds/pod) of 99070 + F, seed from vacuum-emasculated
flowers. With alcohol emasculation, the whole raceme is immersed in a
small beaker of 57% ethyl alcohol for 10 sec, then rinsed in water for a few
seconds. Alcohol is more effective than vacuum for emasculation. How-
ever, alcohol causes more injury to the stigma, so that only about 26% of
the flowers produced pods compared to 60% for vacuum-emasculated
flowers and 76% for nonemasculated controls (Tysdal and Gar!, 1940). It is
recommended that an emasculated, unpollinated control be included when
emasculations are made to determine the effectiveness of procedures and
operators.

C. Pollination

Pollinations can be made at any time of the day or night. Cross-polli-

nation by hand without emasculation is accomplished by alternately trip-
ping flowers of the two parents into a folded cardboard pollen reservoir
(Fig. 3). When parents have normal amounts of pollen and average self-
fertility, about 85% of the seed produced will be hybrid (Pedersen and
Barnes, 1973). By randomly tripping flowers from many plants into the
same pollen reservoir, bulk seed on a population of plants can be produced.

Fig. 3-Tripping alfalfa floret with folded cardboard.

ALFALFA 183

The bench position of plants being pollinated should be changed periodical-

ly to reduce patterns of nonrandom pollination. Viands and Barnes (1977)
estimated that about 3.5 g of seed can be produced on a population of
plants in the greenhouse per person-hour of pollination. The average num-
ber of seed/flower pollinated is about 1.1 from self-pollination and 4.4
from cross-pollination (Pedersen and Barnes, 1973).
Self-pollination requires tripping the flowers without introducing
foreign pollen. The four usual methods of tripping flowers include inserting
a toothpick in the throat of the flower, inserting a toothpick that has the tip
covered with emery paper to rupture the stigmatic membrane, depositing
the pollen into a folded cardboard which retains the pollen for the stigma to
fall into (Fig. 3), and gently rolling or squeezing racemes between the
fingers. Barnes and Stephenson (1971) found that all four methods pro-
duced the same average number of seed per flower tripped. However, roll-
ing racemes was about three times more efficient (pollinations per hour)
than the other three methods. Alcohol should be used to clean hands be-
tween treating plants.

D. Factors Affecting Efficiency

Genetic markers are often used to identify hybrids in pollination

studies. The most useful genetic markers are distinctive ones that can be
identified at the seedling stage, and are not detrimental to plant growth.
About 75 simply inherited traits have been studied in alfafa. The character-
istics of most of these traits have been summarized by Barnes and Hanson
(1967). The best dominant marker available is the red root trait (Clement
and Stanford, 1966). Anthracnose resistance can also be used as a genetic
marker and identified in the seedling stage (Campbell et aI., 1974). The best
recessive markers available are the golden tip trait (Elgin, 1972) and the
green hypocotyl (white flower) trait (Stanford, 1951). Yellow and purple
flower color have been useful genetic markers in many studies; however,
their primary disadvantage is that progenies must be grown to flowering for
classification (Barnes, 1966). All gene markers described in tetraploid alfal-
fa are apparently controlled by tetrasomic inheritance.

V. NATURAL HYBRIDIZATION

Field and cage-planting designs can be important in alfalfa seed pro-

duction because alfalfa, which depends on insects for pollination, varies in
attractiveness to pollinators (Boren et aI., 1962). and produces bot h selfed
and crossed seed. Other factors, such as degree of self-sterility and differ-
ential time and amount of flowering, also influence the extent of crossing
and selfing among parent plants. Seed often is produced by enclosing plants
in a cage with honeybees, leafcutter bees, or bumblebees. Excessive
amounts of selfing (in some cases as much as 900;0) may occur in field cages
because individual honeybees work a single clone instead of randomly visit-
ing all clones (Hanson et a\., 1964). The amount of natural crossing in the
184 BARNES

field between two noninbred alfalfa populations can vary significantly.

Pedersen (1968) reported that 53070 of the seed was hybrid. Kehr (1973) ob-
served differences in crossing from 32 to 96%, with an average of 50070.
Kehr reported that in some situations crossing percentages were significant-
ly higher with an alternate-plant design than with an alternate-row design.
He suggested that alternate plants reduced the possibility of pollinators
being orientated to large blocks of similar plants.
Information on incompatibility and sterility systems in alfalfa was
summarized by Barnes et al. (1972). The gametophytic incompatibility
system only partially prevents self-fertilization. Some plants show virtually
no reduction in fertility after self-fertilization, whereas other plants produce
no selfed seed. There is some evidence for inhibition of pollen-tube growth
within the stigma and style after selfing. The principal expression of self-in-
compatibility appears to be in pollen tube and ovule interactions. Busbice
(1968) proposed that reduced seed yield during inbreeding results primarily
from a loss of heterozygosity in the zygotes which causes lethality.
Both genetic and cytoplasmic-genetic male-sterile systems occur in
alfalfa. The genetic system is characterized by complete degeneration of the
microspores (Childers and McLennan, 1960). The first cytoplasmic male-
sterile system (NS) was described in North America by Davis and Greenblatt
(1967), Bradner and Childers (1968), and Pedersen and Stucker (1969). This
system shows vario!1s degrees of viable pollen production (Barnes and
Garboucheva, 1973; Barnes et aI., 1974). A second cytoplasmic male-sterile
system (ES) has been described in Europe by Guy (1978) and Staszewski and
lakubowska (1978). Genetic steriles have not been used to produce hybrid
alfalfa cultivars. The most efficient pollen control method for hybrid seed
production is cytoplasmic male sterility. A few three-way hybrid cultivars
have been produced utilizing cytoplasmic-genetic male sterility (Barnes et
aI., 1972). Even though the system is workable, the seed production costs
for F, hybrids are higher than those for synthetic cultivars. Seed production
on male-sterile plants must be improved before significant quantities of F,
hybrids can be produced commercially.

VI. SEED DEVELOPMENT, HARVEST, AND STORAGE

Pod formation in alfalfa first becomes evident 2 to 3 days after pollina-

tion. In successful pollinations, the ovary lengthens and extends several
millimeters above the staminal column. In unsuccessful pollinations, the
flowers abscise. Seeds are mature when the pods begin to turn brown. Time
from pollination to seed maturity is 28 days at a constant temperature of 25
C and 40 days at 20 C.
The two methods for harvesting field-grown seed are windrow-curing
and threshing with a pickup combine, or spraying desiccant chemicals and
direct combining (Pedersen et aI., 1972). Greenhouse-produced seed is
usually hand-harvested, air-dried at 35 C, and threshed in some type of
laboratory thresher. Most threshers for small-seeded legumes are custom-
made and are usually designed to rub pods and seeds between two rough
rubber belts or surfaces. One type of thresher for small seed lots was de-
scribed by Norwood (1967).
ALFALFA 185

Hard seeds, those impermeable to water, are common in alfalfa. Ac-

cording to Gunn (1972) the percentage of hard seeds is governed by edaphic
and climatic factors during and after seed maturation and by genetic fac-
tors. Hard-seed percentages of 40 to 50 are common in northwestern-grown
alfalfa, whereas seed lots produced in the southwestern U.S. usually have
less than 20070 hard seed. Greenhouse-grown, hand-harvested seed is often
nearly 100070 hard and should always be scarified. Hard seeds usually are
scarified with custom-made equipment that rubs or polishes the seed against
a rough surface. The seed coat needs to be scratched without injuring the
embryo. Infrared heat treatment (Works and Erickson, 1963) and radio-
frequency electrical treatment (Nelson et aI., 1977) have also successfully re-
duced hard seed content.
Alfalfa seed can be long-lived under certain conditions (Gunn, 1972).
Reports of viable seed after 25 years of storage in dry conditions are not un-
common. When stored under warm, moist conditions, longevity of the seed
will be significantly reduced. Gunn (1972) has suggested the following opti-
mum conditions for long-term storage: fresh, mature seeds with high initial
viability, stored in an environment with less than 10070 moisture (preferably
about 5070), at temperatures near or below 0 C. Low moisture is more im-
portant than low temperature.

VII. TECHNIQUES FOR SPECIAL SITUATIONS

Hormones have not been used in either hybridization research or seed

production. Grafting techniques have been described (Bernardo et aI.,
1970), but have not been used in alfalfa seed research. Embryo culture has
not been highly developed in alfalfa; but it should improve as emphasis on
interspecific hybridization is increased. Recent advances in research with al-
falfa callus tissue culture (Bingham et aI., 1975) indicates a likelihood of
future success in embryo culture research.

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186 BARNES

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ALFALFA 187

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