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24
Measuring Time: Controlling
Development by Photoperiod
and Endogenous Clocks
Two hundred and fifty years ago, Carl v. Linne, better ´
known for his development of the binomial system of
nomenclature, designed a flower clock based on the
opening and closing of the petals at specific but different
times of the day. The plants were arranged in a circle
and one could tell the time of day by simply noting
which flowers were open and which were closed. It is
often difficult for the layman to understand that plants
can tell time without a Timex, but many aspects of
plant behavior can be interpreted in no other way. One
example is the consistent flowering of various species
at particular times of the year. Roses always bloom in
the summer and chrysanthemums in the fall. Indeed,
the flowering of many plants is so predictable from
one year to the next that gardeners have for centuries
incorporated them into their gardens as floral calendars,
unerringly marking the progress of the seasons. In the
northern latitudes, perennial plants sense the short days
of autumn as a signal to induce bud dormancy, thus
anticipating the unfavorable conditions of winter. The
most reliable indication of the advancing season is the
length of day, and an organism’s capacity to measure
the proportion of daylength in a 24-hour period is
known as photoperiodism. However, photoperiodism
is only one of the more outward manifestations of a far
more fundamental timekeeping mechanism, known as
the biological clock.
In this chapter we will examine
• photoperiodism; including the distinction between
short-day plants, long-day plants, and other
response types; the central role of the dark period;
the nature of photoperiodic perception; and a
discussion of current hypotheses to account for the
elusive floral stimulus,
• vernalization—the low-temperature requirement
for flowering in winter annuals and biennial
plants,
• the biological clock, with an emphasis on endogenous rhythms with a 24-hour periodicity and their
role in photoperiodic time measurement,
• the molecular genetic basis for the circadian clock
and the search for the cent