ntensified by screening within the organ.

Pigments,
including but not limited to the photoreceptor itself,
will attenuate the light as it passes through the organ.
Light can also be attenuated by scattering, reflection,
or diffraction within the cells or as it passes between
cells. Thus gradients across individual cells, measured
by using microfiberoptic probes, may vary from 5:1
to 50:1. To further complicate matters, organs such
as coleoptiles appear to function as light pipes. This
means that light applied to the tip, for example, will
be transmitted through the coleoptile to cells further down the organ. Thus the phototropic stimulus
is far from being a simple matter. These complex
interactions between light and the optical properties
of tissue have led to significant difficulties in experimental design as well as in interpretation of the resulting
data.
23.1.2 PHOTOTROPISM IS A
BLUE-LIGHT RESPONSE
Since the 1930s, action spectra for phototropism have
been repeatedly determined for a number of organisms,
but have been most thoroughly documented for coleoptiles of oats (Avena sativa) and maize (Zea mays) and for
sporangiophores of the fungus Phycomyces. The action
spectra for oat coleoptiles and Phycomyces are virtually
identical, indicating that they share a homologous, if not
common, photoreceptor. All other phototropic action
spectra are similar and consistently show two peaks in
the blue region of the spectrum near 475 nm and 450 nm
and a small peak or shoulder at 420 nm (Figure 23.3).
In addition there is a broad action peak in the UV-A
region near 370 nm. The action spectra for both oat and
Phycomyces phototropism show an additional peak in the
region of 280 nm, indicating that the photoreceptor is
probably a chromoprotein.
As early as the 1940s, it was suggested that the
photoreceptor could be a flavin molecule such as riboflavin. However, because of its action spectrum the
photoreceptor for phototropism was subject to the same
flavin-carotenoid controversy described in the previous
chapter for other blue-light responses now known
to be regulated by the cryptochromes. On the other
hand, there were several physiological results that
ruled out carotenoids as the photoreceptor for phototropism long before the responsible pigment, phototropin,
was finally discovered and shown to be a
flavo-protein. Carotenoid biosynthesis, for example,
can be blocked, either by mutation or by treatment
of seedlings with the herbicide norflurazon, which
inhibits the enzyme phytoene desaturase. Yet
carotenoid-deficient maize mutants, albino barley
seedlings, and norflurazon-treated seedlings all exhibit
a normal phototropic response to blue light.
Relative absorption or response
(A)
350 400 450 500
(B)
(C)
Wavelength (nm)
FIGURE 23.3 (A) The action spectrum for Avena coleoptile phototropism. The action spectrum shows peak
activity in the blue and UV-A regions of the spectrum.
For comparison, the absorption spectra for riboflavin, a
common flavonoid (B), and β-carotene (C) are shown.
Although β-carotene ‘‘fits’’ in the blue region (around
450 nm), riboflavin has the required absorption in both
the blue and the UV-A regions (320 nm–400 nm) of the
spectrum.
23.1.3 PHOTOTROPISM ORIENTS
A PLANT FOR OPTIMAL
PHOTOSYNTHESIS
The phototropic blue-light response is distinct from
the blue-light responses mediated by phytochrome
and cryptochrome that were discussed in the previous
chapter. Phytochrome and cryptochrome responses
are morphogenetic responses—they alter the pattern
of growth and development. The singular impact of
phototropism, on the other hand, is that it orients
growth and leaf angle toward incident light in order
to maximize light interception for photosynthesis. The
bending of coleoptiles and hypocotyls is only the most
visible part of a larger blue-light syndrome that plants
use to optimize photosynthesis. Plants also use blue light
to control stomatal opening and facilitate gas exchange
as well as to relocate chloroplasts within the cell.
It has long been known that stomatal opening is
under the control of light. On the one hand, light
absorbed by chlorophyll (i.e., red light) stimulates stomatal opening and obviously depends on photosynthetic
reactions in the guard cell chloroplasts. However, there
is a second, much more sensitive, system that is driven
by low levels of blue light. Most of the evidence points
to a dominant role of the blue-light response in the early
394 Chapter 23 / Tropisms and Nastic Movements: Orienting Plants in Space
phases of stomatal opening, such as when the stomata
open at dawn, prior to the beginning of photosynthesis.
Plants also use blue light to control the high-light
avoidance response of chloroplasts in the mesophyll
cells. In low light, the chloroplasts always gather along
the cell walls that are parallel to the surface, (i.e., periclinal walls) that are perpendicular to the incident light
(Figure 23.4). In high light, such as direct sunlight,
the chloroplasts avoid potential damage by lining up
along the anticlinal walls (i.e., parallel to the incident
light). The redistribution of chloroplasts appears to be
in response to a light gradient through the cytoplasm,
so the responsible photoreceptor is probably located
in the cytoplasm, not the chloroplasts. The mechanism of redistribution has yet to be discovered, but the
cytoskeleton is commonly involved in moving organelles
within the cell and may be involved in the movement of
chloroplasts as well.
23.1.4 FLUENCE RESPONSE CURVES
ILLUSTRATE THE COMPLEXITY
OF PHOTOTROPIC RESPONSES
Perhaps no aspect of phototropism has indicated the
complexity of the process so much as attempts to define
relationships between fluence and response. Phototropism is characterized by a rather curious fluenceresponse
curve, quite unlike most photobiological
responses.
Fluence response curves are generally obtained
by monitoring the response of the organ to different total amounts of light (fluence), usually by using a
single fluence rate but varying the presentation time.
Figure 23.5 shows a fluence-response curve determined
for Avena coleoptile phototropism that illustrates the
classic response to increasing fluence. There is an initial
rise to a first peak, which is called first positive curvature. With increasing fluence, curvature declines, to the
point that this may even result in a bending away from
the light source. This decline and negative response is
called first negative curvature. Note that first negative
curvature is not necessarily ‘‘negative’’ in the sense of
bending away from the light. It may be simply a reduced
positive response. Following the region of first negative
curvature, the response curve again rises into what is
called second positive curvature. In some cases, a second negative and even a third positive curvature have
been reported. First positive curvature is also known
as tip curvature, because it is restricted to the apex of
coleoptiles. Second positive curvature is also called basal
curvature because the curvature extends more toward
the basal region of the coleoptile.
A