.

meristem would be unable to form either petals or

stamens. Not only is this the case, but the whorls that
would normally be filled with petals and stamens are
filled instead with sepal-like and carpel-like structures.
Similarly, in the ag mutant, stamens and carpels are not
formed and all four whorls are filled with sepals or petals.
No doubt this model will continue to be refined as new
genes are discovered and the various genes and their
products are subjected to further molecular analysis.
Although it is becoming more evident which genes
are active in controlling which aspects of floral initiation
and development, their biochemical and physiological function is not yet clear. However, most of the
gene products that have been analyzed thus far share
certain characteristics, called motifs, which are typical of transcription factors. Transcription factors are
DNA-binding proteins that enable RNA polymerases
to recognize promoters and begin transcription of the
gene in eukaryotes. It thus appears that the principal
function of most flowering-time, floral-identity, and
organ-identity genes is to regulate other genes that may
then direct synthesis of the components that actually
make up the individual organs.
25.2 TEMPERATURE CAN ALTER
THE FLOWERING RESPONSE
TO PHOTOPERIOD
There are many examples of interactions between temperature and photoperiod, particularly with respect to
flowering behavior (see Salisbury, 1963, for an extensive listing). In most cases the interaction results in
relatively subtle changes in the length of the critical
438 Chapter 25 / Flowering and Fruit Development
photoperiod or a tendency toward daylength neutrality or an inability to flower altogether at high or low
temperature extremes. There are other plants, however,
for which flowering is either quantitatively or qualitatively dependent on exposure to low temperature. This
phenomenon is known as vernalization. Vernalization
is a means of preventing precocious reproductive development late in the growing season, ensuring instead
that seed production does not begin until the beginning
of the next growing season so that the seed will have
sufficient time to reach maturity.
Vernalization refers specifically to the promotion
of flowering by a period of low-temperature and should
not be confused with other miscellaneous effects of
low-temperature on plant development. The term itself
is a translation of the Russian yarovizatsya; both words
combining the root for spring (Russian, yarov; Latin,
ver) with a suffix meaning ‘‘to make’’ or ‘‘become.’’
Coined by the Russian T. D. Lysenko in the 1920s, vernalization reflects the ability of a cold treatment to make
a winter cereal mimic the behavior of a spring cereal
with respect to its flowering behavior. The response
had actually been observed many years earlier by agriculturalists, but didn’t receive critical attention of the
scientific community until J. G. Gassner showed in 1918
that the cold requirement of winter cereals could be
satisfied during seed germination. For his part, Lysenko
received considerable notoriety for his conviction that
the effect was an inheritable conversion of the winter
strain to a spring strain. His position—a form of the
thoroughly discredited Lamarkian doctrine of inheritance of acquired characteristics—was adopted as Soviet
dogma in biology and remained so until the 1950s.
The adoption of Lysenko’s views as official dogma
had a significant impact on Soviet biology and placed
agriculture in the USSR at a severe disadvantage for
decades.1
25.2.1 VERNALIZATION OCCURS MOST
COMMONLY IN WINTER
ANNUALS AND BIENNIALS
Typical winter annuals are the so-called ‘‘winter’’ cereals (wheat, barley, rye). ‘‘Spring’’ cereals are normally
daylength insensitive. They are planted in the spring
and come to flower and produce grain before the end of
the growing season. Winter strains, however, if planted
in the spring would normally fail to flower or produce mature grain within the span of a normal growing
1The story of vernalization is a classic example of what can
happen when science becomes enmeshed in political
ideology. For the interested student, this unfortunate episode
in the history of science has been artfully documented by
D. Joravsky in his book The Lysenko Affair (Chicago:
University of Chicago Press, 1970).
season. Winter cereals are instead planted in the fall.
They germinate and over-winter as small seedlings,
resume growth in the spring, and are harvested usually
about midsummer. The over-wintering cold treatment,
or vernalization, renders the plants sensitive to long
days.
One of the most thorough studies of vernalization and photoperiodism was carried out on the Petkus
cultivar of rye (Secale cereale) by F. G. Gregory and
O. N. Purvis, beginning in the 1930s. There are two
strains of Petkus rye: a spring strain and a winter strain.
The spring strain of Petkus rye is a facultative long-day
plant. Under short days, floral initiation does not occur
until after about 22 leaves have been produced, typically requiring a season of about 4.5 months. Under
the appropriate long-day regime, however, flowering in
the spring strain is initiated after as few as seven leaves
have been produced, requiring only about two months.
When sown in the spring, the winter strain is insensitive to photoperiod. The winter strain flowers equally
slowly—requiring four to five months—regardless of
daylength.
If seeds of the winter strain are sown in the fall,
however, the germinated seedlings are subjected to
an over wintering low-temperature treatment. When
they resume growth in the spring, winter strain plants
respond as long-day plants in the same way as the spring
strain. The effect of the over wintering cold treatment
can also be achieved by vernalizing the seed, that is, by
holding the germinated seed near 1◦C for several weeks.
Note that the low-temperature treatment, at least in the
case of winter annuals, does not alone promote early
flower initiation. Rather, the effect of vernalization is to
render the seedling sensitive to photoperiod.
Another example of vernalization is seen in biennial plants. Biennials are monocarpic plants that normally
flower (and die) in the second season, again following an over-wintering cold treatment. Typical biennials
include many varieties of sugar- and table-beet
(Beta vulgaris), cabbages and related plants (Brassica
sp.), carrots (Daucus carota) and other members of
the family Umbellifereae, foxglove (Digitalis purpurea),
and some strains of black henbane (Hyoscyamus niger).
Biennials share with the winter annuals the property
that subjecting the growing plant to a cold treatment stimulates a subsequent photoperiodic flowering
response.
Biennials typically grow as a rosette, characterized by shortened internodes, in the first season
(Figure 25.5). Over winter, the leaves die back but
the crown, including the apical meristem, remains
protected. New growth the following spring is characterized by extensive stem elongation, called bolting,
followed by flowering. The cold requirement in
biennials is qualitative (i.e., absolute). In the absence of
a cold treatment many biennials can be maintained in
25.2 Temperature Can Alter the Flowering Response to Photoperiod 439
FIGURE 25.5 Vernalization and stem elongation in cabbage. Left: Cabbage plants were vernalized for six weeks
at 5◦C before being returned to the greenhouse. Center:
Plants were sprayed weekly with a solution containing 5
× 10-4 M gibberellic acid. Right: Control plants grown
at normal greenhouse temperature remain in a rosette
habit. Except for the vernalization treatment, all plants
were maintained in the greenhouse under a long-day
(16-hour) photoperiod.
the nonflowering rosette habit indefinitely. As a rule,
vernalized plants, whether winter annuals or biennials,
tend to respond as long-day flowering plants, although
some biennials are daylength-indifferent following
vernalization. One exception to the rule is the perennial
Chrysanthemum morifolium, a SDP. Some varieties of
Chrysanthemum require vernalization before responding
as a quantitative SDP. As a perennial, Chrysanthemum
normally requires vernalization on an annual basis.
Many other plants such as pea (Pisum sativum) and
spinach (Spinacea oleracea) can be induced to flower
earlier with a cold treatment but it is not an absolute
requirement.
25.2.2 THE EFFECTIVE TEMPERATURE
FOR VERNALIZATION IS
VARIABLE
The range of temperatures effective in vernalization
varies widely depending on the species and duration of
exposure. In Petkus rye, the effective range is -5◦C
to +15◦C, with a broad optimum between +1◦C and
+7◦C. Within these limits, vernalization is proportional
to the duration of treatment. Flowering advances sharply
after as little as one to two weeks’ treatment at 1 ◦C to
2◦C and is maximally effective after about seven weeks
at that temperature (Figure 25.6). Within the effective
range, the temperature optimum is generally higher
for shorter treatment periods. Presumably, a longer
exposure to lower temperatures within the effective
Duration of cold treatment (days)
10 30 50 20 40
40
60
40
20
100
80
Days to flowering
120
FIGURE 25.6 Vernalization in Petkus rye (Secale cereale).
Seeds were germinated in moist sand at 1◦C for the
time indicated. Cold treatments were scheduled so that
all seeds were returned to the greenhouse at the same
time. The number of days to flowering progressively
decreased with increasing length of the cold treatment. (From Purvis, O. N., F. G. Gregory. 1937. Annals
of Botany, N.S. 1:569–591. Copyright, The Annals of
Botany Company.)
range is required because the metabolic reactions leading
to the vernalized state progress more slowly.
Like flowering, the vernalized state is more or less
permanent in most species, giving rise to the concept of an induced state. For example, vernalized
Hyoscyamus, a LDP, can be held under short days
for up to 10 months before losing the capacity to
respond to long-day treatment. On the other hand,
all cold-requiring plants that have been studied are
capable of being devernalized—vernalization can be
reversed if followed immediately by a high-temperature
treatment. Flowering in vernalized winter wheat, for
example, can be fully nullified if the seedlings are held
near 30◦C for three to five days. For most plants, then,
there is a ‘‘neutral’’ temperature where neither vernalization nor devernalization occurs. For Petkus rye the
neutral temperature is about 15◦C. Vernalized seeds of
Petkus rye can also be devernalized by drying them for
several weeks or by maintaining the seeds under anaerobic conditions for a period of time following the col