X 24.

1
HISTORICAL
PERSPECTIVES:
THE DISCOVERY
OF
PHOTOPERIODISM
Although it had earlier been suggested that latitudinal
variations in daylength contributed to plant distribution, the first efforts at controlled experimentation were
conducted by a French scientist in 1912. J. Tournois
found that both Humulus (hops) and Cannabis (hemp)
plants flowered precociously during the winter in the
greenhouse. Tournois eliminated temperature, humidity, and light intensity as environmental cues and in
1914 concluded that the changing of either daylength or
nightlength was responsible for early flowering. Unfortunately, World War I intervened and Tournois did
not live to continue his experiments. At the same time,
H. Klebs was studying flowering of Sempervivum funkii
(commonly known as ‘‘hens-and-chickens’’). Sempervivum grows as a vegetative rosette in the winter
greenhouse. By supplementing normal daylight with
artificial light, Klebs was able to break the rosette habit,
stimulate stem elongation, and induce flowering. From
his experiments, Klebs concluded that length of day
triggered flowering in nature. However, it remained for
W. W. Garner and H. A. Allard to demonstrate the
full impact of daylength on flowering and coin the term
photoperiodism.
W. W. Garner and H. A. Allard were scientists with
the U.S. Department of Agriculture near Washington,
D.C. The initial focus of their work was a mutant
cultivar of tobacco (Nicotiana tabacum), called Maryland
Mammoth. In the field, Maryland Mammoth plants
grew to be very tall with large leaves. Such characteristics
would obviously be advantageous to the tobacco industry
at the time (in the early 1920s), but breeding efforts
were frustrated by the fact that the plants would not
flower in the field during the normal growing season
at that latitude. In the greenhouse, however, even very
small plants flowered in the winter and early spring.
Clearly, flowering was not simply a matter of the age of
the plants. Another problem of interest to Garner and
Allard concerned flowering in soybean (Glycine max).
When the cultivar Biloxi was sown over a 3-month
period from May to August, all of the plants flowered
within a 3-week period in September (Figure 24.1).
The earliest seeded plants thus took 125 days to flower
while those seeded last required only 58 days. Again it
appeared that all plants, regardless of age, were simply
awaiting some signal to initiate flowering.
Like Tournois, Garner and Allard eliminated a
variety of environmental conditions (such as nutrition,
temperature, and light intensity) as the ‘‘signal,’’ coming finally (and with some reluctance) to the conclusion
that flowering was controlled by the relative length of
day and night. Using a crude but effective system of
rolling plant benches in and out of darkened garagelike
buildings at predetermined times, Garner and Allard
proceeded to describe the flowering characteristics of
scores of different species with respect to daylength.
They went on to suggest that bird migration might
also be keyed to daylength—a phenomenon that is now
well documented. We now know that photoperiodic
control is not limited to flowering, but is a basic regulatory component in many aspects of plant and animal
behavior.
May June July August Sept
Days to
Flowering
58
69 F
77 F
92 F
94 FF
125
F
FIGURE 24.1 September soybeans. Soybeans (Glycine
max, cv. Biloxi) sown over a three-month period all
flower within a three-week period in September.
24.1 PHOTOPERIODISM
The switch from the vegetative state to the flowering
state is arguably one of the most dramatic and mysterious
events in the life of a flowering plant. Photoperiodism
influences many aspects of plant development such
as tuber development, leaf fall, and dormancy, but the
control of flowering by photoperiod has attracted the
major share of interest. Indeed, it was the failure of a
tobacco (Nicotiana tobacum) mutant to flower under field
conditions that led to the discovery of photoperiodism
(Box 24.1).
24.1 Photoperiodism 415
24.1.1 PHOTOPERIODIC RESPONSES
MAY BE CHARACTERIZED BY A
VARIETY OF RESPONSE TYPES
Photoperiodic responses fall into one of three general
categories. They are: short-day plants (SD plants),
long-day plants (LD plants), and daylength-indifferent
or day-neutral plants (DNP) (Table 24.1 and
Figure 24.2). Short-day plants are those that flower
only, or flower earlier, in response to daylengths
that are shorter than a certain value within a 24-hour
cycle. Long-day plants respond to daylengths that are
longer than a certain value, while day-neutral plants
flower irrespective of daylength. Within the long- and
short-day categories, we also recognize obligate and
facultative requirements.1 Plants that have an absolute
requirement for a particular photoperiod before they
will flower are considered obligate photoperiodic types.
The common cocklebur (Xanthium strumarium), for
example, is an obligate short-day plant. Xanthium will
not flower unless it receives an appropriate short
photoperiod. On the other hand, most spring cereals
such as wheat (Triticum sp.) and rye (Secale cereale) are
facultative long-day plants. Although spring cereals will
eventually flower even if maintained under continuous
short days, flowering is dramatically accelerated under
long days. The popular research object Arabidopsis
is also a facultative long day plant. However, the
distinction between obligate and facultative response
is not always hard and fast for a particular species or
cultivar. Photoperiod requirement is often modified by
external conditions such as temperature. A particular
species may, for example, have an obligate requirement
at one temperature but respond as a facultative plant at
another temperature.
In addition to these three basic categories, there
are a number of other response types that flower
under some combination of long and short days. Various species of the genus Bryophyllum are, for example,
long-short-day plants (LSD plant)—they will flower
only if a certain number of short days are preceded
by a certain number of long days. The reverse is true
of the short-long-day plant (SLD plant) Trifolium
repens (white clover). A few plants have highly specialized requirements. Intermediate-daylength plants, for
example, flower only in response to daylengths of intermediate length but remain vegetative when the day is
either too long or too short. Another type of behavior
is amphophotoperiodism, illustrated by Madia elegans
(tarweed). In this case, flowering is delayed under intermediate daylength (12 to 14 hours) but occurs rapidly
under daylengths of 8 hours or 18 hours. There are
many, often subtle, variations to the three basic response
1The terms obligate and facultative may be interchanged with
the terms qualitative and quantitative, repectively.
TABLE 24.1 Representative plants exhibiting
the principal photoperiodic response types.
Short-Day Plants
red goosefoot
chrysanthemum
yellow cosmos
poinsettia
Chenopodium rubrum
soybean
Chrysanthemum sp.
tobacco (Maryland Mammoth)
purple perilla
Japanese morning glory
cocklebur
Cosmos sulphureus
Euphorbia pulcherrima
Glycine max
Nicotiana tobacum
Perilla crispa
Pharbitis nil
Xanthium strumarium
Long-Day Plants
Anethum gra