Phototropins and Their LOV Domains：Versatile Plant Blue-Light Receptors

The phototropins phot1 and phot2 are plant blue-light receptors that mediate phototropism, chloroplast movements, stomatal opening, leaf expansion, the rapid Inhibition of hypocotyl growth in etiolated seedlings, and possibly solar tracking by leaves in those species in which It occurs. The phototroplns are plasma membrane-associated hydrophilic proteins with two chromophore domains （designated LOV1 and LOV2 for their resemblance to domains In other signaling proteins that detect light, oxygen, or voltage） in their Nterminal half and a classic serine/threonlne kinase domain in their C-terminal half. Both chromophore domains bind flavin mononucleotide （FMN） and both undergo light-activated formation of a covalent bond between a nearby cystelne and the C（4a） carbon of the FMN to form the signaling state. LOV2-cystelnyl adduct formation leads to the release downstream of a tightly bound amphlpathlc α-helix, a step required for activation of the klnase function. This cysteinyl adduct then slowly decays over a matter of seconds or minutes to return the photoreceptor chromophore modules to their ground state. Functional LOV2 is required for light-activated phosphorylation and for various blue-light responses mediated by the phototroplns. The function of LOV1 is still unknown, although It may serve to modulate the signal generated by LOV2. The LOV domain Is an ancient chromophore module found In a wide range of otherwise unrelated proteins In fungi and prokaryotes, the latter Including cyanobacterla, eubacterla, and archaea. Further general reviews on the phototropins are those by Celaya and Liscum （2005） and Christie and Briggs （2005）.

Light-Regulated Stomatal Aperture in Arabidopsis

The stomatal pores of plant leaves, situated in the epidermis and surrounded by a pair of guard cells, allow CO2 uptake for photosynthesis and water loss through transpiration. Blue light is one of the dominant environmental signals that control stomatal movements in leaves of plants in a natural environment. This blue light response is mediated by blue/UV A light-absorbing phototropins （phots） and cryptochromes （crys）. Red/far-red light-absorbing phytochromes （phys） also play a role in the control of stomatal aperture. The signaling components that link the perception of light signals to the stomatal opening response are largely unknown. This review discusses a few newly discovered nuclear genes, their function with respect to the phot-, cry-, and phy-mediated signal transduction cascades, and possible involve- ment of circadian clock.

New Insights into Dynamic Actin-Based Chloroplast Photorelocation Movement

Chloroplast movement is essential for plants to survive under various environmental light conditions. Photo- tropins--plant-specific blue-light-activated receptor kinases--mediate the response by perceiving light intensity and direction. Recently, novel chloroplast actin （cp-actin） filaments have been identified as playing a pivotal role in the directional chloroplast photorelocation movement. Encouraging progress has recently been made in this field of research through molecular genetics and cell biological analyses. This review describes factors that have been identified as being involved in chloroplast movement and their roles in the regulation of cp-actin filaments, thus providing a basis for reflection on their biochemical activities and functions.

Clues to the signals for chloroplast photorelocation from the lifetimes of accumulation and avoidance responses

Chloroplast photo-relocation movement is crucial for plant survival; however, the mechanism of this phenome- non is still poorly understood. Especially, the signal that goes from photoreceptor to chloroplast is unknown, although the photoreceptors （phototropin 1 and 2） have been identified and an actin structure （chloroplast actin filaments） has been characterized that is specific for chloroplast movement. Here, in gametophytes of the fern Adiantum capillus-veneris, gametophores of the moss Physcomiterella patens, and leaves of the seed plant Arabidopsis thaliana, we sought to characterize the signaling system by measuring the lifetime of the induced response. Chloroplast movements were induced by microbeam irradiation with high-intensity blue light and recorded. The lifetime of the avoidance state was measured as a lag time between switching off the beam and the loss of avoidance behavior, and that of the accumulation state was measured as the duration of accumulation behavior following the extinction of the beam. The lifetime for the avoidance response state is approximately 3-4rain and that for the accumulation response is 19-28 rain. These data suggest that the two responses are based on distinct signals.

FERONIA is involved in phototropin 1-mediated blue light phototropic growth in Arabidopsis

Plant shoot phototropism is triggered by the formation of a light-driven auxin gradient leading to bending growth.The blue light receptor phototropin 1(phot1)senses light direction,but how this leads to auxin gradient formation and growth regulation remains poorly understood.Previous studies have suggested phot1’s role for regulated apoplastic acidification,but its relation to phototropin and hypocotyl phototropism is unclear.Herein,we show that blue light can cause phot1 to interact with and phosphorylate FERONIA(FER),a known cell growth regulator,and trigger downstream phototropic bending growth in Arabidopsis hypocotyls.fer mutants showed defects in phototropic growth,similar to phot1/2 mutant.FER also interacts with and phosphorylates phytochrome kinase substrates,the phot1 downstream substrates.The phot1-FER pathway acts upstream of apoplastic acidification and the auxin gradient formation in hypocotyl under lateral blue light,both of which are critical for phototropic bending growth in hypocotyls.Our study highlights a pivotal role of FER in the phot1-mediated phototropic cell growth regulation in plants.

Blue-light-induced rapid chloroplast de-anchoring in Vallisneria epidermal cells

In the outer periclinal cytoplasm of leaf epidermal cells of an aquatic angiosperm Vallisneria, blue light induces ＂chloroplast de-anchoring＂, a rapid decline in the resistance of chloroplasts against centrifugal force. Chloroplast de- anchoring is known induced within lmin of irradiation with high-fluence-rate blue light specifically, preceding the com- mencement of chloroplasts migration toward the anticlinal cytoplasm. However, its regulatory mechanism has remained elusive, although pharmacological analysis suggested that a calcium release from intracellular calcium stores is necessary for the response, in search of the responsible photoreceptors, immunoblotting analysis using antibodies against phototropins demonstrated that cross-reactive polypeptides of 120-kDa exist in the plasma-membrane fraction prepared from the leaves. In vitro phosphorylation analysis revealed that 120-kDa polypeptides were phosphorylated by exposure to blue light in a fluence-dependent manner. The blue-light-induced phosphorylation activity was sensitive to a Ser/Thr kinase inhibitor, staurosporine, and unusually was retained at a high level for a long time in darkness. Furthermore, phototropin gene homologs （Vallisneria PHOTOTROPINI and PHOTOTROPIN2） expressed in leaves were isolated. We propose that calcium- regulated chloroplast de-anchoring, possibly mediated by phototropins, is an initial process of the blue-light-induced avoidance response of chloroplasts in Vallisneria.

The action of enhancing weak light capture via phototropic growth and chloroplast movement in plants

To cope with fluctuating light conditions,terrestrial plants have evolved precise regulation mechanisms to help optimize light capture and increase photosynthetic efficiency.Upon blue light-triggered autophosphorylation,acti-vated phototropin(PHOT1 and PHOT2)photoreceptors function solely or redundantly to regulate diverse responses,including phototropism,chloroplast movement,stomatal opening,and leaf positioning and flattening in plants.These responses enhance light capture under low-light conditions and avoid photodamage under high-light conditions.NON-PHOTOTROPIC HYPOCOTYL 3(NPH3)and ROOT PHOTOTROPISM 2(RPT2)are signal transducers that function in the PHOT1-and PHOT2-mediated response.NPH3 is required for phototropism,leaf expansion and positioning.RPT2 regulates chloroplast accumulation as well as NPH3-mediated responses.NRL PROTEIN FOR CHLOROPLAST MOVE-MENT 1(NCH1)was recently identified as a PHOT1-interacting protein that functions redundantly with RPT2 to medi-ate chloroplast accumulation.The PHYTOCHROME KINASE SUBSTRATE(PKS)proteins(PKS1,PKS2,and PKS4)interact with PHOT1 and NPH3 and mediate hypocotyl phototropic bending.This review summarizes advances in phototropic growth and chloroplast movement induced by light.We also focus on how crosstalk in signaling between phototro-pism and chloroplast movement enhances weak light capture,providing a basis for future studies aiming to delineate the mechanism of light-trapping plants to improve light-use efficiency.

Phototropism in land plants： Molecules and mechanism from light perception to response

BACKGROUND： Phototropism is the response a plant exhibits when it is faced with a directional blue light stimulus. Though a seemingly simple differential cell elongation response within a responding tissue that results in organ curvature, phototropism is regulated through a complex set of signal perception and transduction events that move from the plasma membrane to the nucleus. In nature phototropism is one of several plant responses that have evolved to optimize photosynthesis and growth. OBJECTIVE： In the present work we will review the state of the field with respect to the molecules and mechanisms associated with phototropism in land plants. METHODS： A systematic literature search was done to identify relevant advances in the field. Though we tried to focus on literature within the past decade （1998-present）, we have discussed and cited older literature where appropriate because of context or its significant impact to the present field. Several previous review articles are also cited where appropriate and readers should seek those out. RESULTS： A total of 199 articles are cited that fulfill the criteria listed above. CONCLUSIONS： Though important numerous and significant advances have been made in our understanding of the molecular, biochemical, cell biological and physiologic mechanisms underlying phototropism in land plants over the past decade, there are many remaining unanswered questions. The future is indeed bright for researchers in the field and we look forward to the next decade worth of discoveries.