hot1, while second positive curvature is mediated by

both phot1 and phot2. Phot1 and photo2 contribute

equally to stomatal opening, while the avoidance movement of chloroplasts under high light intensities is
mediated only by phot2.
Recent evidence has also indicated a role for
phototropins in the promotion of cotyledon and leaf
expansion and the rhythmic sleep movements of kidney
bean leaves (see Chapter 24).
23.1.7 A HYBRID RED/BLUE LIGHT
PHOTORECEPTOR HAS BEEN
ISOLATED FROM A FERN
A particularly interesting photoreceptor has recently
been isolated from the fern Adiantum capillus-veneris.
Designated neochrome (formerly known as phy3),
this photoreceptor has properties of both phytochrome
and phototropin (Figure 23.7). The amino acid
sequence of the amino-terminal domain shows a
significant homology to the chromophore-binding
domain of phytochrome. Furthermore, when the gene
was expressed in yeast and the purified protein was
reconstituted with a phycocyanobilin chromophore, it
showed typical phytochrome photoreversible behavior.
But neochrome also has the two LOV domains, which
bind FMN, and a serine/threonine kinase domain
at the C-terminus that are virtually identical to
phototropin. Neochrome is required for phototropism
in Adiantum, which is regulated by red as well as blue
light. Neochrome is clearly a hybrid photoreceptor
that mediates red, far-red, and blue-light responses.
What is curious, however, is that Adiantum also has
two fully functional phototropins like their higher
plant counterparts and, again like their higher plant
counterparts, phot2 is solely responsible for mediating
high-light chloroplast avoidance movements.
Several other ferns, mosses, and algae have both
phototropins and neochromes, which raises interesting
questions regarding the evolution of photoreceptors as
well as their physiological action.
23.1.8 PHOTOTROPIN ACTIVITY
AND SIGNAL CHAIN
Participants in the phototropin signal chain are only
now just beginning to ‘‘come to light.’’ Some of the
more important factors can be summarized as follows:
1. Autophosphorylation of phototropin plays a significant
role in the phototropic response, probably by initiating
a phosphorylation cascade. Studies employing mutants
of the PHOT1 gene have shown that the protein is
apparently folded in such a way that the phosphorylation site is blocked by the LOV2 domain in the
dark. Absorption of blue light by the chromophore
induces a change in the conformation of the protein so that the phosphorylation site is available and
active. The role of the LOV1 domain is not clear.
Mutants lacking the LOV1 domain have shown that
LOV1 is not necessary for phosphorylation but its
presence does increase kinase activity.
2. Phototropins may be involved in gene regulation. No
substrates directly phosphorylated by phototropin
in planta have yet been identified, but there are
several proteins that interact with the photoreceptor and are necessary for a proper response.
For example, the proteins NONPHOTOTROPIC
HYPOCOTYL 3 (NPH3) from Arabidopsis and a
homologous protein (called an ortholog) from rice,
COLEOPTILE PHOTOTROPISM 1 (CPT1),
include domains that are characteristic of transcriptional regulators or proteins that are involved in
protein degradation. The mutants nph3 and cpt1,
in which these proteins are missing, show no phototropic response.
3. Phototropin disrupts polar auxin transport. One of the
challenges presented by phototropism is to establish
whether or not a link exists between the absorption
398 Chapter 23 / Tropisms and Nastic Movements: Orienting Plants in Space
of blue light by phototropin and the asymmetrical
auxin distribution proposed by the Cholodny-Went
hypothesis. As described previously, asymmetric
auxin distribution has been demonstrated experimentally. Recent experiments have focused on the
relationship between phototropin and auxin efflux
facilitator PIN1. PIN1 is normally localized at the
basal ends of xylem-associated cells where it
serves to facilitate the polar vertical flow of auxin
(Chapter 18). When the location of PIN1 in Arabidopsis hypocotyls was monitored by immunofluorescence
microscopy following phototropic stimulus,
the basal location of PIN1 in wildtype plants
was disrupted in the cortical cells on the shaded
side of the hypocotyl. A similar disruption was
not observed in phot1 mutants. These results
suggest that phototropic bending is initiated by
a phototropin-mediated decrease in the vertical
transport of auxin. This would lead to a retention
or sequestering of auxin, and consequent increased
growth, in those cells that are directly involved in
phototropic bending.
23.1.9 PHOTOTROPISM IN GREEN
PLANTS IS NOT WELL
UNDERSTOOD
A final area of concern is phototropism in light-grown
plants, where relatively little is known about the
phototropic process. As with phytochrome, discussed in
Chapter 22, most of what we know about phototropism
is derived from laboratory studies with etiolated seedlings. However, in light-grown cucumber (Cucumis
sativus) and sunflower (Helianthus annuus) seedlings
subjected to uniform lighting, curvature of the
stem can be induced by simply shading one of the
FIGURE 23.8 Curvature in cucumber (Cucumis sativus)
seedlings induced by shading cotyledons. The left-hand
cotyledon was covered with aluminum foil and the
seedling uniformly irradiated with white light for 8
hours. (After Shuttleworth, J. E., M. Black. 1977. Planta
135:51.)
cotyledons (Figure 23.8), and the phototropic response
of sunflower seedlings is markedly decreased if the
leaves are removed. The cucumber response, at least,
differs from the classical phototropic response in that
it is induced by red light rather than blue light. This
appears to be a phytochrome-mediated response and is
related to inhibition of hypocotyl elongation below the
irradiated cotyledon. Both the cucumber and sunflower
responses may be attributed to the fact that the leaves
are a prime source of auxin required for the growth
response. Both cucumber and white mustard (Sinapis
alba) also exhibit a classical phototropic response
induced by irradiating the hypocotyls directly with blue
light. Clearly the control of stem growth in green plants
is an area where there is still much to learn.
23.2 GRAVITROPISM
Gravitropism is probably one of the most unfailingly
obvious and familiar plant phenomena to most people
(Figure 23.9). Everyone is aware that shoots always
grow ‘‘up’’ and roots always grow ‘‘down.’’ Or do they?
A casual walk through the woods or garden should
reveal how overly simplified this view is. The lateral
branches of most trees and shrubs do not grow up; they
grow outward in a more or less horizontal position.
Stolons (or runners) of stra