OsPIN1a Gene Participates in Regulating Negative Phototropism of Rice Roots

The complete open reading frame of OsPINla was amplified through reverse transcriptase- polymerase chain reaction （RT-PCR） based on the sequence deposited in GenBank to explore the relationship between the auxin efflux protein OsPINla and the negative phototropism of rice roots. Sequencing results showed that the GC content of OsPINla was 65.49%. The fusion expression vector pCAMBIA-1301-OsP/N1a：：GFP containing the OsPINla gene and a coding green fluorescent protein （gfp） gene was constructed. The fusion vector was transferred into onion epidermal cells by Agrobacterium tumefaciens transformation. The transient expression of OsPINla-GFP was mainly located in the nucleus and cell membrane. Moreover, the transgenic plants were obtained by Agrobacterium-mediated genetic transformation. Molecular detection performed by using PCR and β-glucuronidase staining showed that the target construct was integrated into the genome of rice. The negative phototropic curvatures of the transgenic rice roots were higher than those of the wild type. Similarly, the expression levels of OsPINla in the transgenic plants were considerably higher than those in the wild-type plants. These results suggest that OsPINla is crucial in the negative phototropic curvature of rice roots.

Effect of Indoleacetic acid (IAA) on the Negative Phototropism of Rice Root

To explore the effects of IAA on negative phototropism of rice ( Oryza sativa L.) root, agar block containing IAA was unilaterally applied on root tip to examine the phototropic response of root to exogenous IAA, and microstructure of the bending part was observed with an optical microscope. The growth of seminal roots could be regulated by exogenous IAA as well as light, as a result the root bent towards the site treated, causing asymmetric growth of the root cells at the elongation zone and consequently bending growth. IAA concentration in the shaded side of adventitious root increased much greater at 1.5 h after the start of irradiation. The unequal lateral IAA distribution can be concluded to be the main cause for negative phototropism of rice root.

Calcium Signaling is Involved in Negative Phototropism of Rice Seminal Roots

Calcium ions （Ca2＋） act as an intracellular second messenger and affect nearly all aspects of cellular life. They are functioned by interacting with polar auxin transport, and the negative phototropism of plant roots is caused by the transport of auxin from the irradiated side to the shaded side of the roots. To clarify the role of calcium signaling in the modulation of rice root negative phototropism, as well as the relationship between polar auxin transport and calcium signaling, calcium signaling reagents were used to treat rice seminal roots which were cultivated in hydroculture and unilaterally illuminated at an intensity of 100-200 pmol/（m2.s） for 24 h. Negative phototropism curvature and growth rate of rice roots were both promoted by exogenous CaCI2 lower than 100 pmol/L, but inhibited by calcium channel blockers （verapamil and LaCI3）, calcineurin inhibitor （chlorpromazine, CPZ）, and polar auxin transport inhibitor （N-l-naphthylphthalamic acid, NPA）. Roots stopped growing and negative phototropism disappeared when the concentrations increased to 100 pmol/L verapamil, 12.500 ~Jmol/L LaCI3, 60 pmol/L CPZ, and 6 pmol/L NPA. Moreover, 100 pmol/L CaCI2 could relieve the inhibition of LaCI3, verapamil and NPA. The enhanced negative phototropism curvature was caused by the transportation of more auxin from the irradiated side to the shaded side in the presence of exogenous Ca2＋. Calcium signaling plays a key role as a second messenger in the process of light signal regulation of rice root growth and negative phototropism.

The Phototropism of Jurassic Petrified Wood in North China Plate

Trees on the side directly exposed to sunlight generally grow faster than on the opposite side, a phenomenon termed plant phototropism. There are in situ vertical trunks of silicified wood in the Xiadelongwan area of Yanqing County, north Beijing, where the first National Geologic Park of Petrified Wood of China has been built since 2002. A few trunks have well-preserved growth rings. One petrified stump from the formation shows a positive phototropism direction of SW230°. As compared with the modern normal growth stumps in Beijing plain area, which have a positive phototropism direction of SW210 °± 5°, the evidence of wood phototropism supports the conclusion of previous palaeomagnetic studies that the North China Plate has rotated clockwise since the Late Jurassic. The known petrified wood stumps in the Yanshan-Liaoning area are mainly found from the strata of 165-136 Ma, which corresponds to the main stage of the Yanshanian Movement.

Phototropism of Petrified Wood and Its Relation with the Rotation of Different Blocks in China and the Possibility of Application in the World

Normally, trees on the side directly exposed to sunlight will grow faster than the opposing side. This phenomenon is termed plant phototropism. Moreover, palaeomagnetists have revealed that the Junnar Block has never rotated since the Mesozoic. The petrified woods in the Jiangjunmiao area of Qitai County show the positive phototropism direction of SSW220. By compared with the modern normal growth stumps in plain area, which have positive phototropism direction of SSW 219 ± 5, this observation supports the conclusion of palaeomagnetic researchers: the Junggar basin has never rotated since the Late Jurassic.

Phototropism in the Marine Red Macroalga Pyropia yezoensis

Phototropism is a response to the direction of light that guides growth orientation and determines the shape of plants to optimize photosynthetic activity. The phototropic response is present not only in terrestrial plants but also in water-living algae. However, knowledge about phototropism in Bangiophycean seaweeds is limited. Here, we examined the phototropic response of the red alga Pyropia yezoensis to elucidate the regulatory mechanism of phototropism in Bangiophyceae. When leafy gametophytes and filamentous sporophytes of P. yezoensis were cultured under directional light, phototropism was observed in the gametophytes. Conchosporangia on the sporophytes also exhibited phototropism. Phototropism was positive in the majority of gametophytes and conchosporangia but in some cases was negative. In addition, a strong phototropic response occurred under white light, whereas blue and red light elicited minor and no responses, respectively. This observation is in contrast with the phototropic response in terrestrial plants and several algae, in which blue light is responsible for positive phototropism. Surprisingly, the genome of P. yezoensis has no homologues of the photoreceptors for blue and red light, revealing differences in the regulation of phototropism between terrestrial plants and P. yezoensis . Studies on the phototropism in P. yezoensis could shed light on the evolutional divergence of phototropic responses in plants.

Negative phototropism of rice root and its influencing factors

Some characteristics of the rice (Oryza sativa L.) root were found in the experiment of unilaterally irradiating the roots which were planted in water: (ⅰ) All the seminal roots, adventitious roots and their branched roots bent away from light, and their curvatures ranged from 25° to 60°. The curvature of adventitious root of the higher node was often larger than that of the lower node, and even larger than that of the seminal root. (ⅱ) The negative phototropic bending of the rice root was mainly due to the larger growth increment of root-tip cells of the irradiated side compared with that of the shaded side. (ⅲ) Root cap was the site of light perception. If root cap was shaded while the root was irradiated the root showed no negative phototropism, and the root lost the characteristic of negative phototropism when root cap was divested. Rice root could resume the characteristic of negative phototropism when the new root cap grew up, if the original cells of root cap were well protected while root cap was divested. (ⅳ) The growth increment and curvature of rice root were both influenced by light intensity. Within the range of 0-100μmol@m-2@s-1, the increasing of light intensity resulted in the decreasing of the growth increment and the increasing of the curvature of rice root. (ⅴ) The growth increment and the curvature reached the maximum at 30℃ with the temperature treatment of 10-40℃. (ⅵ) Blue-violet light could prominently induce the negative phototropism of rice root, while red light had no such effect. (ⅶ) The auxin (IAA) in the solution, as a very prominent influencing factor, inhibited the growth, the negative phototropism and the gravitropism of rice root when the concentration of IAA increased. The response of negative phototropism of rice root disappeared when the concentration of IAA was above 10 mg@L-1.

Phototropism of Thalli and Rhizoids Developed from the Thallus Segments of Bryopsis hypnoides Lamouroux

Newly regenerated thalli were used to study the phototropism of Bryopsis hypnoides Lamouroux under different qualities of light. Positive phototropism in the thalli and negative phototropiam In the rhizoida of B. hypnoides were investigated and analyzed in terms of bending. Both thaiii and rhlzoids developed from thallus segments exhibited typical tip growth, and their photoreceptive sites for phototroplam were also restricted to the apical hemisphere. The bending curvature of rhizoids and thalli were determined with unilateral lights at various wavelengths and different fluence rates after a fixed duration of Illumination. The trends of bending from the rhizoid and thallus were coincident, which showed that the action spectrum had a large range, from ultraviolet radiation （366.5 nm） to green light （524 nm）. Based on the bending curvatures, blue light had the highest efficiency, while the efficiency of longer wavelengths （〉500 nm） was significantly lower. External Ca^2＋ had no effect on the bending curvature of thalli and rhlzolda. Blue light （440 nm） induced thallus branching from rhizoids, while red light （650 nm） had no such effect. Fast-occurring chloroplast accumulation In the outermost cytoplasmic layer of the blue light （440 nm）-Irradiated region In the rhizoid was observed, from which protrusions （new thalli） arose after 4 h of the onset of illumination, and this action was thought to be driven by the dynamics of actin microfilamenta.

Negative Phototropism of Chlorophytum comosum Roots and Their Mechanisms

The aerial roots of Chlorophytum comosum were grown hydroponically,allowing us to study the performance and mechanism of negative phototropism. The results of this study were as follows. All the adventitious roots and their branch roots bent away from light with a maximum curvature of approximately 88.5°. Blue-violet light prominently induced negative phototropism while red light had no effect. The root cap was the site of photo perception. Roots with shaded or divested root caps exposed to unilateral light showed no negative phototropism,but resumed their original characteristics when the shade was removed or when new root caps grew. The curvature increased when the light intensity ranged 0–110 μmol·m^(-2)·s^(-1). The negative phototropism curvature could be promoted by exogenous CaCl_2 but was inhibited by exogenous LaCl_3; exogenous CaCl_2 could reduce the inhibitory effect of LaCl_3. Unilateral light induced the horizontal transport of IAA from the irradiated side to the shaded side,resulting in an unequal distribution of IAA in both the sides,leading to negative phototropism. The horizontal transport of IAA was promoted by exogenous Ca^(2+) but inhibited by exogenous La^(3+).

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.

Positive/Negative Phototropism:Controllable Molecular Actuators with Different Bending Behavior

Herein,a series of molecular actuators based on the crystals of(E)-2-(4-fluorostyryl)benzo[d]oxazole(BOAF4),(E)-2-(2,4-difluorostyryl)benzo[d]oxazole(BOAF24),(E)-2-(4-fluorostyryl)benzo[d]thiazole(BTAF4),and(E)-2-(2,4-difluorostyryl)benzo[d]thiazole(BTAF24)showed unique bending behavior under UV irradiation.The one-dimensional(1D)crystals of BOAF4 and BTAF4 bent toward light,whereas those of BOAF24 and BTAF24 bent away from light.Although the chemical structures of these compounds are similar,the authors found that F···H–C interaction played a key role in the different molecular packing in structures crystals,which led to the positive/negative phototropism of the actuators.Moreover,theoretical calculations were carried out to reveal the mechanical properties of the crystals.Taking advantage of these photomechanical properties,the authors achieved the potential application in pushing objects,as well as enriching and removing pollutants.Hence,the molecular actuators with different bending behavior could be fabricated by introducing different number of F atom,which may open a novel gate for crystal engineering.

The photosensory function of Zmphot1 differs from that of Atphot1 due to the C-terminus of Zmphot1 during phototropic response

The role of phot1 in triggering hypocotyl phototropism and optimizing growth orientation has been wellcharacterized in Arabidopsis, whereas the role of Zmphot1 in maize remains largely unclear. Here, we show that Zmphot1 is involved in blue light-induced phototropism. Compared with Atphot1, Zmphot1exhibited a weaker phototropic response to very low-fluence rates of blue light(< 0.01 μmol m-2s-1),but stronger phototropic response to high-fluence rates of blue light(> 10 μmol m-2s-1) than Atphot1. Notably, blue light exposure induced Zmphot1-green fluorescent protein(GFP), but not Atphot1-GFP, to form the aggregates in the cytoplasm of Nicotiana benthamiana cells. Furthermore, by generating the chimeric phot1 proteins, we found that the serine-threonine kinase(STK) domain at the C-terminus is responsible for a more volatile membrane association of Zmphot1. Consistently, the chimeric phot1 protein fusing the STK domain of Zmphot1 with other domains of Atphot1 responded similarly as Zmphot1 to both low and high fluence rates of blue light. Interestingly, although both Zmphot1 and Atphot1 interact with AtNPH3, Zmphot1 induced weaker dephosphorylation of NONPHOTOTROPIC HYPOCOTYL 3(NPH3) than Atphot1. Together, our findings indicate that Zmphot1 and Atphot1 exhibit different photosensory function during phototropic response and that the STK domain may play a key role in determining their properties.

Associative learning in plants:light quality history may matter

The possibility of associative learning in plants is a topic of ongoing controversy.In one published study,growing pea plants were reported to associate two stimuli(airflow and light)and thereafter use one(airflow)as an indicator for the other(light),similar to dogs in Pavlov’s famous experiments.However,this observation could not be independently repeated.Here we examine a possible reason for the failure of a published reproduction attempt,which used substantially different light quality during plant cultivation prior to experimental treatments than in the original study.This could have resulted in dramatically different growth characteristics.While the relevance of the original report of plant associative learning remains questionable,greater attention should be paid to good documenting and standardizing the light conditions,in particular spectral quality,not only in studies of plant learning and memory,but also in other areas of experimental plant biology.

A Botanist’s Cognitive View on Plant Growth: Cross-Talk between Developmental and Sensitivity Networks

An alteration in plant phenotypes assisted by their responses to the environmental stimuli (=tropism) has been fundamental to understand the “plant sensitivity ” that plays a crucial role in plants’ adaptive success. Plants succeed through the deployment of moderators controlling polar auxin-transport determining organ bending. Stimulus-specific effectors can be synthesized by the outer peripheral cells at the bending sites where they target highly conserved cellular processes and potentially persuade the plant sensitivity at large. Remarkably, the peripheral cells require different time-intervals to achieve the threshold expression-levels of stimulus-specific molecular responders. After stimulus perception, tropic curvatures (especially at growing root-apices) are duly coordinated via integrated chemical and electrical signalling which is the key to cellular communications. Thus, the acquired phenotypic alterations are the perplexed outcome of plant’s developmental pace, complemented by the sensitivity. A novel aspect of this study is to advance our understanding of plant developmental-programming and the extent of plant-sensitivity, determining the plant growth and their future applications.

Seven Things We Think We Know about Auxin Transport

Polar transport of the phytohormone auxin and the establishment of localized auxin maxima regulate em- bryonic development, stem cell maintenance, root and shoot architecture, and tropic growth responses. The past decade has been marked by dramatic progress in efforts to elucidate the complex mechanisms by which auxin transport regulates plant growth. As the understanding of auxin transport regulation has been increasingly elaborated, it has become clear that this process is involved in almost all plant growth and environmental responses in some way. However, we still lack information about some basic aspects of this fundamental regulatory mechanism. In this review, we present what we know （or what we think we know） and what we do not know about seven auxin-regulated processes. We discuss the role of auxin transport in gravitropism in primary and lateral roots, phototropism, shoot branching, leaf expansion, and venation. We also discuss the auxin reflux/fountain model at the root tip, flavonoid modulation of auxin transport processes, and outstanding aspects of post-translational regulation of auxin transporters. This discussion is not meant to be exhaustive, but highlights areas in which generally held assumptions require more substantive validation.

Photoreceptor-Mediated Bending towards UV-B in Arabidopsis

Plants reorient their growth towards light to optimize photosynthetic light capture--a process known as phototropism. Phototropins are the photoreceptors essential for phototropic growth towards blue and ultraviolet-A （UV- A） light. Here we detail a phototropic response towards UV-B in etiolated Arabidopsis seedlings. We report that early differential growth is mediated by phototropins but clear phototropic bending to UV-B is maintained in photl phot2 double mutants. We further show that this phototropin-independent phototropic response to UV-B requires the UVoB photoreceptor UVR8. Broad UV-B-mediated repression of auxin-responsive genes suggests that UVR8 regulates directional bending by affecting auxin signaling. Kinetic analysis shows that UVR8-dependent directional bending occurs later than the phototropin response. We conclude that plants may use the full short-wavelength spectrum of sunlight to efficiently reorient photosynthetic tissue with incoming light.

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）.

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.

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.