Circadian Rhythms The Clock Within

Organisms are normally subjected to daily cycles of light and darkness, and both plants and animals often exhibit rhythmic behavior in association with these changes. Examples of such rhythms include leaf and petal movements (day and night positions), stomatal opening and closing, growth and sporulation patterns in fungi (e.g., Pilobolus and Neurospora), time of day of pupal emergence (the fruit fly Drosophila), and activity cycles in rodents, as well as metabolic processes such as photosynthetic capacity and respiration rate.

When organisms are transferred from daily light-dark cycles to continuous darkness (or continuous dim light), many of these rhythms continue to be expressed, at least for several days. Under such uniform conditions the period of the rhythm is then close to 24 hours, and consequently the term circadian rhythm is applied (see Chapter 17). Because they continue in a constant light or dark environment, these circadian rhythms cannot be direct responses to the presence or absence of light but must be based on an internal pacemaker, often called an endogenous oscillator. A molecular model for a plant endogenous oscillator was described in Chapter 17.

The endogenous oscillator is coupled to a variety of physiological processes, such as leaf movement or photosynthesis, and it maintains the rhythm. For this reason the m (m

Flowering stage:

Vegetative stage: ■

Oldest plant

(6-7 leaves),

flowering after

1 LD cycle^

Younger plant

(4-5 leaves),

flowering after

2 LD cycles^

/

Youngest plant

(2-3 leaves), \

flowering after

4 LD cycles^------

i i i

Number of LD cycles

Number of LD cycles endogenous oscillator can be considered the clock mechanism, and the physiological functions that are being regulated, such as leaf movements or photosynthesis, are sometimes referred to as the hands of the clock.

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