VERNALIZATION AND THE GROWTH-PHASE

CONCEPT
H. H. McKINNEY
Bureau of Plant Industry
INTRODUCTION

The term vernalization is the anglicized form of the Russian

word iarovization. It signifies that certain winter annuals and bien-
nials can be induced to follow the spring-annual habit by suitable
treatment of the germinated seed or the active bulbs before planting,
thus making it possible to obtain sexual reproduction the first season
from spritig plantings. However, as the term is now used by some
workers it embraces practically all of the environmental factors and
all the methods applied at any time in the plant's development and
which are capable of accelerating sexual reproduction in any species
of plant. As the term iarovization was first used it referred to
treatments which altered certain germinated seeds physiologically,
these alterations in turn changing the direction of growth in suit-
able subsequent environments. The term was not applied to the
breaking of a rest period as in the potato and many seeds. How-
ever, there has been a tendency among some workers to apply the
term vernalization in this connection. In fact, it has been applied
by Shchernetsky (63) to the process of soaking sugar beet seed in
water for 4 days at 17.2 ~ to 21.1~ C. before planting with the inten-
tion of increasing the tonnage and sugar content of the roots.
In general it may be stated that the fundamental concept or prin-
ciples of vernalization are based on the following facts: (1) that
species and varieties possess different optimum environmental re-
quirements during their several critical growth phases, (2) that
these optimum requirements must be met within certain limits,
otherwise sexual reproduction will not occur or it will be delayed,
and (3) that certain of these environmental requirements which are
naturally supplied during the first developmental phases in certain
plants can be supplied artificially to the slightly germinated seeds or
to bulbs before planting.

ItISTORICAL BACKGROUND

In the present article reference will be made to those papers

which appear the most pertinent, and no attempt will be made to
26
26 THE BOTANICAL REVIEW

cite all of the many titles in the literature. However, an attempt

is made to cite papers which have good literature lists. These taken
together with the present list should enable any student to become
acquainted with the entire field very easily. Papers having rather
large literature lists are so indicated in the literature list of this
paper.
During recent years, Lysenko (42, 43, 44, 46), a Russian inves-
tigator, has devoted much time to the study of vernalization and
growth phases in many crop plants. In Russia this line of physiol-
ogy is being carried forward on a very large scale; however, it
seems only fair to point out that the basic concepts (54, 70, 71) in-
volved in these studies have been known within certain circles for
many years. They simply have not been recognized in all circles
of plant science until recently.
The older horticultural and agricultural journals and the older
scientific journals and text books show that some growers and some
botanists recognized part or all of these concepts. To take the
specific phenomenon, vernalization of wheat, on which Lysenko
developed his views, it is found that in 1857 Klippart (35), an early
observer and close student of crops who was associated with the
Ohio State Board of Agriculture, published a rat.her clear elucida-
tion of this phenomenon in one of his annual reports to the State
Board. However, 20 years previous to Klippart's record it was
recorded that a grower in New York had produced a crop of grain
from spring-sown winter-wheat seed which had been subjected to
low temperatures before seeding. In 1850 Allen (1), an agricul-
tural expert of that day, recorded a similar observation. It appears,
therefore, that in Klippart's time the idea of vernalization was more
than local in the United States. Since Klippart's record is so clear
and to the point it is quoted as follows:
"To convert winter into spring wheat, nothing more is necessary
than that the winter wheat should be allowed to germinate slightly
in the fall or winter, but kept from vegetation by a low temperature
or freezing, until it can be sown in the spring. This is usually done
by soaking and sprouting the seed, and freezing it while in this
state and keeping it frozen until the season for spring sowing has
arrived. Only two things seem requisite, germination and freezing.
It is probable that winter wheat sown in the fall, so late as only to
germinate in the earth, without coming up, would produce a grain
which would be a spring wheat, if sown in April instead of Septem-
ber. The experiment of converting winter wheat into spring wheat
 VERNALIZATION 27

has met with great success. It retains many of its primitive winter
wheat qualities, and produces at the rate of 28 bushels per acre."
In 1883 Hellriegal (26) concluded that barley has a lower opti-
mum temperature during the tiller-formation phase than during the
stem-elongation phase.
In 1893 the phenologists, ecologists and horticulturists met with
the International Meteorological Congres~ in Chicago and their
several papers were published by the U. F~. Department of Agricul-
ture (68). At that meeting Paul Schreiber from Chemnitz, Ger-
many, presented an ecological traper in ",vhich he stated :
"The growth of vegetation requires heat, water and sunshine ; but
of each the proper measure, as every excess or deficiency acts in-
juriously. It should, therefore, he the object of our investigations
to determine how much of heat, water and sunshine is required by
different plants, and how these influential factors are to be dis-
tributed during the various phases of plant life.
"Our information concerning the duration and power of sunshine
is increasing so rapidly that we may hope for early and important
additions to our knowledge concerning these elements of our inves-
tigations. If our labors in this direction are to be of practical value
to the husbandman, they must include careful notations of the suc-
cessive phases of plant life or, at least, of the main phases of growth
--the so-called phenologic observations. If, in this manner, we
discover the laws governing each separate phase or phenomenon,
and from them the joint result of their reciprocal influences, our
object will have been accomplished."
W. Detmer of Jena, Germany, presented a physiological paper
in which he makes the following statement:
"The position of the three cardinal points, the minimum, opti-
mum, and maximum, is by no means the same for the various
physiological processes which take place in a plant or an organ; it
also varies for a given process in different species of plants, and is
even influenced by the degree of development of an organ."
Dr. Egon Ihne of Friedberg, Germany, made the following state-
ment in his paper on plant phenology :
"Although the same vegetal phase may set in on a date varying
from year to year, the date depending primarily on the climate of
each year, yet to reach this phase the plant requires an amount of
heat that is constant from year to year. A plant may, therefore, be
considered as a means for measuring heat; and the beginning of a
certain vegetal phase is also a standard for measuring a certain sum
total of heat supplied up to that date, and this sum total expresses
28 THE BOTANICAL REVIEW

the measure of heat required by the plant to reach ~ phase in ques-

tion.
"Further, as has been demonstrated by physiological /nvestiga-
tions, it is not every temperature above zero (v.entigrade) that is
effective in vegetation, but the degree at which the temperature
begins to be effective varies for different plants and phases. In
the simple addition of positive temperatures, be they shade means,
shade maxima, or insolation maxima, this vaviatfffity finds no ex-
pression. The zero point of effective temperature should first be
determined for each phase and plant in question."
Klebs (34) in 1918 emphasized the importance of recognizing the
different requirements of plants during the several phases of de-
velopment. In oCen~pervic,u m Fttnckii he distinguished three dis-
tinct phases, each having different environmental requirements.
The same year Gassner (17) reported that earliness in winter
wheat is favored when low temperatures obtain during the seedling
stage and high temperatures obtain during the later stages of
development.
In 1923 (48) the writer called attention to a field test in which a
good crop of winter wheat was obtained in Illinois from seed sown
late in November but which did not emerge until spring. In an-
other paper McKinney and Sando (52) deal with additional litera-
ture on the subject.
From the records cited there can be no doubt that the funda-
mental concepts relating to vernalization and to growth-phase
requirements are of long standing in America and in Europe.

METHODS

The initial steps in vernalization may be summarized as follows:

Seeds are first soaked in a suitable quantity of water to just start
germination, after which they are subjected to suitable temperatures
for periods ranging from about 5 to 60 days, depending on the
species and variety. Or in the case of some plants as beets, cab-
bage, cauliflower, carrots and turnips, the young seedlings are sub-
jected to low temperatures for suitable periods.
In general there are two ranges of temperature in use, low tem-
peratures ranging from slightly above freezing to about 10 ~ C. and
high temperatures ranging from 20 ~ to 30* C. It is claimed that
darkness is an essential fcature during exposure in the high-tem-
perature method, but darkness is of no importance in the low-
temperature treatment of germinated wheat (52) nor in the high-
temperature treatment of germinated corn (64).
 VERNALIZATION 29

In all cases germination is started but retarded by holding the

moisture content of the seed at suitable percentages ranging from
about 45% to 60% of the dry weight of the seed, depending on
the species. In some cases salts have been added to the water to
prevent excessive germination during the treatment (5). At the
completion of the treatment, seeds may be dried for a short time
before sowing or they may be sown in a slightly moist condition.
The low-temperature treatment is used for the seeds of wheat,
barley, rye, oats, timothy, rice, meadow foxtail, vetch, rape, lentils,
white lupines, crimson clover, red clover, Austrian winter field peas,
carrots, beets, turnips, and cabbage, and it has been applied in some
modified form to such bulbs as those of the Easter lily, daffodil,
Dutch and Spanish iris.
The high-temperature treatment has been advocated for corn,
cotton, soybeans, millet sorghum, Sudan grass, and rice.

RESULTS

In this review it seems unnecessary to consider the results of

more than a few experiments which bring out principles. In the
cereal plants, emphasis will be placed on wheat and rye because
these plants have been given the most intensive study and because
the principles established in the studies of these plants seem to
apply to other winter annual members of the Gramineae in the tribes
Hordeae and Aveneae.

Cereal Plants
LOW TEMPERATURE PROCESS
In all types of wheat tested thus far by the writer and Sando (50,
52), sexual reproduction is not dependent on a critical temperature
or a critical photoperiod, because this process occurs over very wide
ranges of both factors. However, the time when sexual reproduc-
tion occurs and also the yield of seed are influenced by temperatures
and the photoperiod (32, 33, 52) and by light intensity (52).
Furthermore, the optimum conditions for the earliest sexual repro-
duction are not the same as the optimum conditions for the highest
seed yield (S).
When winter wheats such as Harvest Queen and Turkey are
grown at high temperatures in a long day throughout the life cycle,
they produce many side shoots or tillers, each of which has many
internodes and Ieaves; the elongation of the internodes, head forma-
30 THE BOTANICAL REVIEW

tion, and sexual reproduction are retarded. On the other hand,

when the seed is germinated slightly and chilled at temperatures
slightly above freezing for 60 days before it is sown at high tem-
peratures in a long day, the resulting plants behave as spring wheats.
They develop relatively few tillers which produce few internodes
and leaves, the internodes elongate rapidly and heading and sexual
reproduction are accelerated. In other words, the chilling treat-
ment has not broken a dormant period such as occurs in perennial
temperate-zone fruits, but it has stimulated a directional change in
the plant's activities when growth continues in a suitable environ-
ment. On the one hand, the plant vegetates excessively and sexual
reproduction is delayed, whereas on the other hand vegetation is
reduced and sexual reproduction is accelerated.
So far as they have been studied, winter barley (6, 7), winter
oats (6, 8) and winter rye (59) have given ample evidence of react-
ing essentially in the same manner indicated for wheat.
Some varieties such as Purplestraw wheat are intermediate or
facultative with respect to seasonal growth habit. They require a
moderate amount of low temperature during early growth for early
sexual reproduction. Some late spring wheats such as Kinney will
head earlier if low temperatures are supplied during early growth.
In order to determine the commercial value of vernalization in
the United States field trials were carried out with winter and
spring wheats at Rosslyn, Va. ; Lincoln and Alliance, Nebraska;
Hays, Kansas; Mandan and Langdon, North Dakota; and Mocca-
sin, Montana (51).
The most satisfactory yields were obtained at Langdon, North
Dakota, the most northerly station. At that station Kanred winter
wheat from seed chilled 50 days outyielded all the unchilled Mar-
quis spring-wheat controls, though the difference was slight.
Slightly greater yields were obtained from certain chilled spring
wheats than from the unchilled spring-wheat controls, but the same
spring varieties at Mandan gave the highest yields when the seed
was not chilled.
Although rapid acceleration of sexual reproduction reduces the
yield of seed per plant because of a reduction in the number of
tillers and in the number of fertile florets per head (52), yields have
been higher when the plants were cultured under suitable short days
and low growing temperatures during the initial growth phase (50,
 VERNALIZATION 31

52) than when the germinated seeds were vernalized in the usual
manner.
Harvest Queen plants from germinated seeds vernalized for 40
days near 33 ~ F. and grown with uninterrupted light at summer
temperatures, headed 89~ days from the time the seed was put to
soak (50). This is the most rapid heading time yet obtained in
Harvest Queen winter wheat from chilled germinated seed. How-
ever, only 10 to 30 seeds per plant were produced.
Yields of 75 seeds per plant have been obtained in Harvest Queen
when heading took place 88 days from the time the seed started to
soak and when the seedlings and plants were grown according to the
following schedule of temperatures, photoperiods, and times (50) :
21.1 ~ to 23.9 ~ C. on moist filter paper for two days to start ger-
mination.
10.0 ~ C. for 36 days with a photoperiod of 8 hours in culture
chamber.
15.6 ~ C. for 18 days with a photoperiod of 8 hours in culture
chamber.
21.1 ~ to 23.9 ~ C. to end of test with a photoperiod of 18 hours in
greenhouse.
These plants actually headed one day earlier than those from
vernalized seed, and it is likely that still earlier heading is possible
without reducing the yield to 30 seeds per plant.
Harvest Queen plants from chilled germinated seeds will yield
75 seeds if grown at 21.1 ~ to 23.9 ~ C. with a daily photoperiod of
16 to 18 hours. However, 100 or more days are required for plants
to head when computing time from the beginning of the soaking
process (50).
Although varieties of winter wheat differ in their temperature
and length-of-day requirements for earliness (50), tests indicate
that all the winter varieties tested complete their life cycles quite
rapidly and produce good yields of seeds when grown near 7.2 ~ to
10.0 ~ C. with a daily photoperiod of 8 to 10 hours for 45 days, fol-
lowed by temperatures near 21.1 ~ to 23.9 ~ C. with a daily photo-
period of 16 to 18 hours. Exposure of 20 days to the low tempera-
tures and short days has been satisfactory for, the facultative
varieties such as Purplestraw.
In hybridizing work Sando and the writer have obtained essen-
tially simultaneous flowering in early, intermediate and late varieties
32 TttE BOTANICAL REVIEW

of both winter and spring wheats, thus making possible all com-
bination crosses.
In addition to hastening maturity it is claimed by Russian
workers that vernalization increases drought resistance in spring
cereals, and it is now claimed that drought resistance is increased
when the seeds are moistened and dried intermittently three times
at moderate tempet:atures, 20 ~ to 22 ~ C., allowing germination to
proceed slowly during the moist stages (27, 28, 29, 30, 57, 58).
Germination is started by adding water amounting to 30 per cent
of the dry weight of the seed. The seed is then dried, remoistened
with water amounting to 20% of the dry weight of seed, dried
again, remoistened with water amounting to 15~ of dry weight of
seed, dried and sown. The object is to allow very slight progress
of germination at each moistening and to keep the plumules and
roots from developing to a point which prevents easy driUing. The
method is referred to as prehardening.
Data cited by Zuhr (77) indicate that vernalization reduced bunt
and loose smut in a spring variety and in a facultative variety of
wheat grown in the field. Loose smut was reduced 45.3% to
68.3% in comparison with non-vernalized controls.
These aspects of the vernalization problem have not been tested
to any extent in other parts of the world.
Vasiljev (69), Timofeeva (66) and others found that winter
hardiness in winter wheats is reduced by vernalization. It appears
that vernalization of the germinated seed before sowing prevents
the normal hardening which takes place in the field during the
autumn and early winter.
Builina (9) found that the optimum time for vernalization tends
to increase with the degree of winter hardiness characteristic of a
given variety of winter wheat. The varieties having low winter-
hardiness required 45 to 55 days, those of medium hardiness re-
quired 55 to 60 days, whereas the most winterhardy required 60 to
65 days.
Rice was vernalized at 3 ~ to 5 ~ C. by Ossewaarde (55) in Hol-
land. The time periods were 2 weeks and 5 weeks. The vegeta-
tive period was shortened 2 to 7 days and the yield of straw and
grain were increased by the two-weeks treatment.
The high-temperature method has been used also for vernalizing
rice as indicated below.
 VERNALIZATION 33

HIGH-TEMPERATURE PROCESS

It has been claimed that certain summer annuals have a short

vegetative period when the slightly germinated seeds are held for 5
to 15 days at temperature ranging from 20 ~ to 30 ~ C. in darkness
(45).
Seed of several cereal crop plants has been treated according to
the methods recommended and comparisons have been made in the
field between the plants from the treated and the nontreated seed.
Sprague (64) working with several hybrids, inbreds and one
variety of corn Zea mays gave the method a rather thorough trial
at the Arlington Experiment Farm.
Although the seed was soaked in a 0.5% solution of Uspulun to
cut down molds, vernalization reduced the germination greatly over
the controls and caused delay in emergence in most cases.
Vernalization reduced the number of internodes very slightly and
hastened sexual maturity. However, the differences in time of
maturity between the vernalized and the control series were so slight
as to be of no agronomic importance. Yields of grain were reduced
in the vernalized series. In the five comparisons which were sig-
nificant the reduction was approximately 15% to 30%. As in the
case of wheat the day length following vernalization influences the
time of maturity of the corn plant. Sixteen hours of light daily
during culture hastened sexual maturity from 10 to 15 days over
continuous light. Vernalization was just as effective in continuous
light as it was in the dark.
Tests with vernalized corn were conducted by Elema (15) in
Holland and the results agree in general with those reported by
Sprague.
Martin reports (53) that seed of 41 varieties of sorghum were
vernalized by the recommended high-temperature method and all
varieties failed to mature earlier than the controls. In most
varieties vernalization greatly reduced or completely destroyed
germination. Similar results have been reported from Holland by
Elema (15) and van Hock (31).
Haigh conducted tests with rice in Ceylon, and his results were
reported in a personal communication to the Imperial Bureau of
Plant Genetics (2). Seed was soaked for 24 hours, then held for
6 days at threedifferent temperatures, 25 ~ 30 ~ and 35 ~ C. Sow-
ings were made in the field. Vernalization favored flowering by
34 THE BOTANICAL REVIEW

5 to 10 days in comparison with the dry-seed controls, and by 1 to

4 days in comparison with the controls sown with germinated, non-
vernalized seed. The method has no practical value, according to
the report.

Forage Plants
~epikova (11, 12) reports that Alopecurus pratensis (meadow
foxtail) and Phleum pratense (timothy) produced seed the first
season when the germinated seeds were first vernalized near 3 ~ C.
Trifolium pratense (red clover, double-cut type) produced seed the
first season when the seed was vernalized at 10~ to 12~ C. for 10
days.
Zerling and ~epikova (75, 76) carried out additional tests and
found that Trifolium pratense (red clover, single-cut type) seeded
the first season when vernalized 20 to 40 days at 3 ~ to 8 ~ C. These
investigators report that vernalized meadow foxtail, timothy and
red clover were earlier and gave higher yields than the controls the
second season. In other words, the autumn, winter and spring
conditions seemingly did not have an equalizing effect on the con-
trols when compared with the vernalized lots. The timothy seemed
to show the greatest benefit the second season from vernalization
of the year previous.
Kostov (38) reported favorable results with vetch vernalized for
10 days at 8 ~ to 10~ C.
2vanskii (73) and 2vanskii and Spilevskii (74) report that seed
was obtained from winter rape the first season when the germinated
seeds were kept in snow for 16 days and in a cool cellar for 15 days
before sowing on April 23.
Soybean seeds were vernalized by the high-temperature method
in Holland by Elema (15) and by van Hoek (31) and in England
at the Long Ashton Station (2, 118). All reports indicate that
plants from vernalized seed behaved as the plants from the non-
vernalized seed. Negative results were reported from Long Ash-
ton with respect to peas vernalized by the low-temperature method.
In the United States McKee (47) tested the effect of low-tem-
perature vernalization on White lupine (Lupinus albus), crimson
clover (Trifolium incarnatum) hairy vetch (Vicia villosa), Aus-
trian Winter field pea (Pisum arvense), double-cut red clover (T.
pratenae), and white sweet clover (Melilotus alba). Germinated
 VERNALIZATION 35

seeds were held at 0 ~ C. for 40 days before sowing. All species

except the red clover and sweet clover came into flower and fruit
when vernalized. Nonvernalized lots either did not bloom or were
late. The temperatures were probably too low and the time too
long in the case of the clovers.
The same investigator tested foxtail millet (Chctetachloa italica),
Sudan grass (Sorghum vulgare sudanense ), soybean ( Soja max)
and crotalaria by the high-temperature method of vernalization.
In most cases vernalization decreased vigor and in no case did the
treatment hasten the time of maturity.

Miscellaneous Plants
Burr and Turner (10) vernalized seeds of tomato and cucumber
at 1 ~ to 3 ~ C. for 7 to 44 days before sowing. Vernalization re-
duced germination, stunted the plants and reduced fruit yields. In
later tests these investigators ( T u r n e r and Burr 67) found that
vernalization at - 0 . 2 8 ~ C. followed by continuous illumination of
the seedling and young plants for 2 to 24 days hastened maturity
and increased the yield of fruit in tomato. The failure of the first
test is attributed to the fact that the vernalized seed was kept too
long before planting.
Yamamato (72) reports that seed production is favored in radish
(Raphanus sativus L.) when the small seedlings are first subjected
to 0 ~ to 5 ~ C. for 10 to 15 days before placing at warm tempera-
tures.
Van Hoek (31) vernalized potato tubers in a lighted box at 18 ~
to 20 ~ C. for 26 days. Plants from vernalized tubers grew more
rapidly, were more vigorous and matured 6 days sooner than the
controls. Twenty-three hills from vernalized and from nonver-
nalized tubers yielded 22.3 and 20.6 kg tubers, respectively. Tubers
from the vernalized series were larger than those from the control.
Chesnokov (13) tested beet seeds (Beta vulgaris var. Egyptian
and Bordeaux) vernalized 43 days and 68 days at 3 ~ to 5 ~ C. The
longer time of exposure increased the amount of bolting the first
season. This treatment induced bolting in almost 2/3 of the plants
in both varieties and seed was produced. Similar results have been
reported by others (4).
In southern New Mexico (56), high yields of sugar beet seed
are obtained in late June and July from seed sown in the field dur-
36 THE BOTANICAL REVIEW

ing late August or early September. Here vernalization takes place

naturally during the mild winter.
In a later paper Chesnokov (14) states that 80% to 90% of the
beet plants bolted the first season when the young seedlings were
chilled for 50 days. H e claims turnip, cabbage and carrot seeded
more freely the first season when the young seedlings were chilled
than when slightly germinated seeds were chilled.
Ta~lanov and Pudovkina (65) vernalized germinated cotton
seeds at 13 ~ to 23 ~ C. for 3 days followed by 25 ~ to 30 ~ C. for 13
days. Vernalization hastened flowering and maturity 9 to 11 days
in the Egyptian varieties; acceleration was especially evident in the
late varieties. The American varieties showed less acceleration
and more variations between themselves than was the case in the
Egyptian types. It is claimed that the yields of all Egyptian varie-
ties and part of the American varieties were increased. Tests car-
ried out with cotton in Indore (2, 134) led to the conclusion that
vernalization has no commercial advantage in that locality.
The acceleration of blooming in ornamental plants has been prac-
ticed for many years by growers and by amateurs. A recent book
by Lawrie and Poech (40) covers this field rather fully. Some of
the methods used involve the breaking of a true rest period previous
to the final treatments which accelerate blooming. High and low
temperatures during storage and after potting or planting are used.
The acceleration of blooming in daffodil and bulbous iris was
studied by Griffiths (25). He found that daffodil bulbs should be
kept under ordinary shed storage conditions. I f temperatures go
below 21 ~ C. additional heat should be supplied. When the buds
are visible--about August 1--they are stored at 10 ~ C. for about
one month, then potted and held at 10 ~ to 18 ~ C. for about a month
in a cellar. If the bulbs are then benched in a greenhouse with
night temperatures gradually increasing from 10 ~ to 15 ~ C., flower-
ing begins just before Christmas, depending on the variety.
Early blooming in Dutch and Spanish iris is accomplished by
storing at 26.7 ~ C. from the time of digging to August 1. After
this treatment they are stored at 10 ~ C. for a month, then planted
in a well ventilated greenhouse. Blooming occurs in December
and January, depending on the variety.
In Bermuda (3) growers and experimenters accelerate blooming
in the Easter lily when bulbs are stored at 2.22 ~ C. for a month
before planting October 15.
 VERNALIZATION 37

MORPHOLOGICAL AND PHYSIOLOGICAL STUDIES

McKinney and Sando (50) reported that earliness in wheat is

correlated rather closely with a number of leaves and internodes
produced by the culms or stalks. A number of early, intermediate
and late spring varieties were studied with different daily photo-
periods in two ranges of temperature. Early varieties have fewer
leaves and internodes than late ones, also temperature and light
conditions favoring few leaves and internodes in a given variety
tend to favor earliness except that change in time of flowering is
somewhat more sensitive to environmental change than is the inter-
node number. For example, Prelude headed from 32 to 43 days
after planting with a leaf number of 5. When conditions induced
11 leaves per culm, heading was extended to 140 days. The variety
Reward headed from 33 to 53 days from planting with a leaf num-
ber of 6. When 13 leaves per culm were produced, heading was
extended to 104 days.
Under conditions favoring the earliest heading--continuous light
at summer temperatures--the earliest varieties produced 5 leaves
and the latest produced 11 leaves per culm.
Purvis (59) working with winter rye found that earliness is
favored by the conditions which favor few leaves per culm. The
same results were also observed and recorded by McKinney and
Sando (52) for winter wheat. Using Petkus spring and winter
rye Purvis and Gregory (60) found that approximately the first
seven of the lateral primordia of the main axis are obligate leaf
primordia, the subsequent 18 lateral prlmordia are labile or faculta-
tire in that they may become leaves or spikelets, depending on the
temperature and photoperiod, and the subsequent ones are obligate
spikelet primordia.
Harvest Queen winter wheat (52) has approximately seven obli-
gate leaf primordia and approximately fifteen labile primordia.
These numbers are inherently less in the early spring and winter
wheats and greater in the late varieties.
The number of stalks per plant (tiUering) (6, 16, 32, 59) and
the number of seeds per plant are intimately connected with earli-
ness. The writer and Sando (50) reported that few tillers and
small numbers of seeds result when winter wheat is forced to com-
plete its life cycle too rapidly, but that seed yields are higher when
forcing is less rapid and is accomplished by means of low growing
38 THE BOTANICAL REVIEW

temperatures with short days, followed by the higher temperatures

and long days, than is the case when the germinated seeds are ver-
nalized before seeding and the subsequent plants are forced by high
temperatures and long days.
The writer with Sando (52) found that immature germinated
seeds of Harvest Queen winter wheat vernalized as efficiently as
mature germinated seeds in a refrigerator. However, they did not
test seeds earlier than the soft-dough stage. Kostjucenko and
Zarubailo (36) report that the maturing seeds of winter wheat are
vernalized naturally in the field in northern areas when the tem-
peratures are sufficiently low before the seed is mature. Gregory
and Purvis (19, 22) report the same phenomenon in winter rye but
their tests with wheat failed. They found that winter-rye heads
chilled during the middle period of ripening vernalized, whereas
those chilled before and after this period did not. They (18, 22,
24) also succeeded in vernalizing excised germinated winter-rye
embryos on nutrient agar containing carbohydrate at 1o C. In later
tests Gregory and de Ropp (24) found excised embryos grown on
nutrient agar containing no carbohydrate failed to vernalize whereas
those on agar containing 3% sucrose vernalized.
Kostju~enko and Zarubailo (37) in summarizing their work con-
clude that the milk-ripe stage responds to vernalization because the
physiology of the seed at this time is nearly comparable to the seed
during germination. They cite data relating to the carbohydrate
content, peroxidase and eatalase activities in support of their con-
clusion. These investigators conducted field tests in northern and
more southern areas in Russia and found that natural vernalization
during maturity in the north shortened the vegetative period the fol-
lowing season when the seed was sown at more southern stations in
comparison with plants from seed grown at the southern station the
previous season. They point out that natural vernalization must
be taken into account when seed of winter and late spring cereals
is taken from northern to southern areas, otherwise genetic and
agronomic results may not be properly interpreted. They indicate
that winter hardiness is greatly reduced by natural vernalization
and they recommend that seed of winter cereals intended for fall
sowing in north Russia come from the more southern regions where
natural vernalization does not occur. Temperatures at 15 ~ C. and
below during the milk stage of the seed are regarded as favoring
natural vernalization.
 VERNALIZATION 39

In the United States, the daily mean temperature may reach 15 ~

C. during the midperiod of seed development at some points near
the Canadian border. However, practically no winter wheat is
grown in that latitude and it is still a question as to the vernalization
response in the Durum wheats which are later than most of the
common spring varieties.
In their studies on winter rye Gregory and Purvis (20) found
that devernalization can take place. They ran a test in 5 parts, each
for a period of 6 weeks. Germinated seeds were subjected to 1 ~
C. and to 20 ~ C. in darkness for alternating periods. The 1 ~ C.
treatments were for 1, 2, and 3 and 6 day intervals, respectively.
After each of these intervals there was in each case exposure to 20 ~
C. for 1 day, followed by the respective schedules above at 1 ~ C.
While at 1 ~ C. the seedlings h a d access to the ordinary or normal
atmosphere but while at 20 ~ C. the seeds were in an atmosphere of
nitrogen. A control received 1 ~ C. every day, but the nitrogen and
ordinary atmosphere were alternated daily. After the treatments
the seeds were planted and cultured at suitable growing tempera-
tures and photoperiods. The seed lots receiving the 1-degree treat-
ments for 1, 2, 3 and 6 consecutive days and the control seed lot
receiving 1~ C. continuously had 0, 20, 60, 100 and 100 per cent of
the seeds vernalized, respectively. It is concluded that high tem-
perature nullifies the effect of low temperature.
In later tests these investigators also (21, 23) found that an
atmosphere of nitrogen extended the vegetative period in spring rye.
Rye seeds thus devernalized were revernalized when subjected to
1 ~ C. for 3 weeks in the normal atmosphere.
Winter rye (22) which had been vernalized and then dever-
nalized lost its ability to head early at high temperatures, but it
produced more tillers at high temperatures than rye which had
never been vernalized.
From the results cited it is to be expected that warm day tem-
peratures will nullify the natural vernalization induced in maturing
seeds by low night temperatures until a sufficiently low daily mean
temperature is reached. The exact relative efficiencies of low
temperatures for vernalization and of high temperatures for dever-
nalization when these phenomena are working against each other
apparently has not been determined in terms of time and degrees of
temperature.
40 THE BOTANICAL REVIEW

Germination, though progressed very slightly, and at least 50~5

moisture are essential for the successful vernalization of mature
seeds (41, 43, 44, 52), when the temperature is at optimum.
Darkness and light had no apparent influence on the vernaliza-
tion efficiency of low temperatures in the case of germinated winter-
wheat seed (52).
Turkey winter wheat seed was held at 3.3 ~ C. for 61 days. One
sample was in total darkness during the test and another sample
received daylight during the entire day each day of the test. After
vernalization the seeds were sown outdoors during early summer
with the natural photoperiod. The plants from seed chilled in day-
light headed 47 days after sowing whereas the plants from seed
chilled in darkness headed 49 days after sowing. A difference of
two days is not significant thus indicating that low temperatures
and not darkness stimulated the early heading. Turkey wheat
sown during the summer without vernalization does not head in the
vicinity of Washington, D. C.
Whyte (70) indicates that these findings are inconsistent with
earlier findings published by McKinney and Sando (49, 50, 52).
However, the writer fails to find such inconsistency. The earlier
tests referred to relate to plants growing at temperatures consider-
ably above 38 ~ F. and in short days vs. long days, whereas the light
and dark test referred to above was carried out at 3.3 ~ C. with ger-
minated seeds, essentially the conditions of seed vernalization, and
the conclusion relates only to slightly germinated seed (vernaliza-
tion), a point which seems clear enough from the chapter headings
and descriptions in that paper (52, 630).
Tests and observations reported by the writer and Sando (52)
ruled out the endosperm, the tips of the roots, coleoptile tip and tip
of the first true leaf of seedlings as the sole active centers of
sensitivity to the vernalization process.
Krasnoseljskaja-Maximova (39) claimed that spring cereals con-
tain no detectable substance which favors early flowering but that
the winter cereals contain an inhibiting substance which must be
counteracted before the plants can proceed to sexual reproduction.
Later Sereiskii and Sluckaja (62) claimed that winter wheat con-
tains no such inhibitor.
Richter (61) in his report as director summarizes the results
obtained in studies conducted under the direction of Cailahjan on
vernalization and photoperiodism as follows:
 VERNALIZATION 41

Vernalization shifted the iso-electric point of albumino-lipoids

towards the acid end, increased the permeability of the protoplasm
and the mobility of the albuminous complex, intensified photosyn-
thesis, increased dry matter in insoluble proteins, and decreased
soluble protein. It is claimed that sexual processes controlled by
light occur in the leaves and are related to the formation of flower
hormones (floregin) which moves to the promeristem. In grafting
experiments the floregin was transferred from stock to scion. Flo-
regin was found not to be specific for species or biologic forms. It
is claimed that no substance inhibiting or retarding flowering is
formed in the leaves.
Purvis and Gregory (22) have shown from studies with excised
embryos and with ripening grain that vernalization in winter rye is
localized in the embryo and is independent of changes in the endo-
sperm or aleurone layer. They conclude that the growing embryo
is able to synthesize hormones at low temperatures from a substrate
containing glucose, and inorganic salts including nitrates. It is
their idea that a precursor (A) in the embryo of winter rye is con-
verted by autocatalysis at low temperatures into a substance (B)
which in turn may be converted into a spikelet initiating substance
(C) and a spikelet maturation substance (D) or into a vegetative
leaf-promoting substance (E), depending on the subsequent photo-
period or temperature, and the system C ~:~ B is reversible. Dever-
nalization due to drying is accounted for by a conversion of sub-
stance (B) to substance (E). Substance (B) is naturally in high
concentration in spring varieties.

DISCUSSION
In general the chilling method of vernalization has been found
to accelerate sexual reproduction with greater certainty than the
high-temperature method in the particular species for which each
method has been recommended. Many workers report that they
have been unable to obtain acceleration with the high-temperature
method and in those cases where acceleration has occurred the com-
mercial advantage has not been evident.
For several years vernalization methods have been tested in many
parts of the world and it is noteworthy that the majority of investi-
gators outside of Russia fail to recognize any great commercial
value to be derived from the methods as applied to the small grains,
rice, corn, sorghum, forage crops and cotton in the regions where
42 THE BOTANICAL REVIEW

these crops are adapted. It seems to be the general consensus of

opinion that the crop problems can best be solved through develop-
ing better adapted genotypes.
Some commercial value is attached to the chilling method when
used to force flowering in daffodils, Dutch and Spanish iris, and
Easter lily. The method seems to offer commercial possibilities for
speeding up seed production in certain biennials such as the garden
beet and sugar beet, and it has considerable value in speeding up
seed production in genetic and plant-improvement work with many
crop plants. However, entirely aside from the stimulating influ-
ence of low temperatures during germination, it is strikingly evident
that the temperature, the photoperiod, the intensity and the quality
of light 'during the entire period of growth have marked influence
on the time when sexual reproduction occurs and it is only when
these factors are understood in relation to all the growth phases of
the particular genotype that the most satisfacory results can be
obtained from the initial chilling.
In view of the evidence, it seems justifiable to conclude that Har-
vest Queen, Turkey and similar winter wheats and other winter
cereals are not typical long-day plants with respect to their earliest
sexual reproduction, but are what may be termed short-day --> long-
day plants, and they may be considered as low-temperature --> high-
temperature plants. This method of expression indicates that low
temperature or short days and low temperatures in combination
must obtain in the germinating seeds or in the young growing
plants, respectively, for suitable periods in order that the first labile
primordium and subsequent ones will develop into spikelets instead
of leaves when the high temperatures and the long days are intro-
duced. A similar situation seems to apply to the facultative
varieties and to certain late varieties commonly placed in the spring
group, but in these the initial optimum temperatures are either not
so low or the periods of exposure to low temperatures are shorter
than is the case with the strictly winter varieties.
The typical spring cereals are high-temperature and long-day
plants in the truest sense with respect to early sexual reproduction.
However, as in the case of winter varieties the spring varieties
differ with respect to their optimal environmental requirements for
the earliest completion of the life cycle.
Earliness of sexual reproduction appears to depend on the inter-
relation of several plant characters, i.e., (1) characteristic number
 VERNALIZATION 43

of obligate leaf primordia and attending internodes, (2) character-

istics of the embryo which bring into existence or activate sub-
stances which in turn activate the labile lateral primordia into spike-
lets at characteristic (a) temperatures, (b) photoperiods and (c)
periods of time, (3) growth characteristics of the stem internodes
and (4) rate of maturity of the sex organs and seeds.
Conditions favoring the earliest sexual reproduction induce rela-
tively low yields of seed per plant. In Marquis spring wheat and
Harvest Queen winter wheat high seed yields obtain when five to
seven of the labile primordia produce leaves and when the heads on
3 to 4 stalks have 14 to 15 well-filled spikelets. This is accomplished
by gradual changes from the lower to the higher growing tempera-
tures and from the short to the longer days. The total time
requirement is greater than required for the earliest reproduction.
On the basis of the reversals in the order of heading in certain
pairs of varieties when grown in different environments, it can be
concluded that segregating populations from certain parent crosses
will not give constant segregating ratios for earliness under all
conditions of temperature and day length. As pointed out in a
previous paper (50), populations which are segregating for earli-
ness and lateness should be tested and classified as far as practicable
under several conditions of temperature and photoperiod to facili-
tate the selection of genotypes homozygous for the several charac-
ters influencing earliness and lateness.
These conclusions do not vitiate the fundamental postulates of
evolution or of genetics but they do indicate that a knowledge of
growth phases and character expression in relation to environmental
factors will facilitate the adequate planning of many genetic studies
and the interpretation of the results. Such characters as yield, sea-
sonal growth habit, recumbence, relative earliness, resistance and
susceptibility to certain parasites, to extreme temperatures and to
drought, serve to illustrate a few of the complexes which can be
studied profitably under several conditions of environment for the
purpose of determining at least some of the simpler characters of
which they are composed. In critical genetic studies these simpler
characters offer advantages over the complex ones.
The responses of the healthy or the diseased plant to temperature,
the photoperiod, light intensity, light quality, humidity, soil mois-
ture, soil fertility, etc. are most certainly determined by internal
mechanisms which in turn constitute genetic characters, and
44 Tile BOTANICAL ggVIEW

although these ~ r a e t e r s cannot be m e a s u r e d d i r e c t l y a t p r e s e n t , in

s o m e cases t h e y can b e m e a s u r e d i n d i r e c t l y a n d b e dealt w i t h as
,-haracters. A s this p r o c e d u r e is followed, a n d as it becomes m o r e
generally recognized t h a t dominance a n d segregation r a t i o s in m a n y
c h a r a c t e r s m u s t be considered within well defined environmental
limits, some o f the c o n f u s i o n r e g a r d i n g the e n v i r o n m e n t a n d
inheritance will disappear.
T h e v a r i o u s e x p e r i m e n t a l results discussed in this p a p e r seem to
shed no direct light on L y s e n k o ' s 1 view that certain g e n o t y p e s can
be altered b y systematic culture in an e n v i r o n m e n t which differs
f r o m that n a t u r a l l y required by the plant. T h i s so-called alteration
of the g e n o t y p e - - r e f e r r e d to as " t r a i n i n g " the p l a n t - - w o u l d vir-
tually a m o u n t to directed m u t a t i o n on a m a s s - p r o d u c t i o n scale. T o
settle this point it would be necessary to plan a n d conduct the ex-
p e r i m e n t s in a somewhat different m a n n e r than was followed b y
the w r i t e r a n d the other investigators cited in the vernalization a n d
g r o w t h - p h a s e studies.

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 VERNALIZATION 47

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