Bacteriophytochromes are photochromic histidine kinases using a biliverdin chromophore.

Phytochromes comprise a principal family of red/far-red light sensors in plants. Although phytochromes were thought originally to be confined to photosynthetic organisms, we have recently detected phytochrome-like proteins in two heterotrophic eubacteria, Deinococcus radiodurans and Pseudomonas aeruginosa. Here we show that these form part of a widespread family of bacteriophytochromes (BphPs) with homology to two-component sensor histidine kinases. Whereas plant phytochromes use phytochromobilin as the chromophore, BphPs assemble with biliverdin, an immediate breakdown product of haem, to generate photochromic kinases that are modulated by red and far-red light. In some cases, a unique haem oxygenase responsible for the synthesis of biliverdin is part of the BphP operon. Co-expression of this oxygenase with a BphP apoprotein and a haem source is sufficient to assemble holo-BphP in vivo. Both their presence in many diverse bacteria and their simplified assembly with biliverdin suggest that BphPs are the progenitors of phytochrome-type photoreceptors.     

Phytochromes and light signal perception by plants--an emerging synthesis

For plants, the sensing of light in the environment is as important as vision is for animals. Fluctuations in light can be crucial to competition and survival. One way plants sense light is through the phytochromes, a small family of diverse photochromic protein photoreceptors whose origins have been traced to the photosynthetic prokaryotes. During their evolution, the phytochromes have acquired sophisticated mechanisms to monitor light. Recent advances in understanding the molecular mechanisms of phytochromes and their significance to evolutionary biology make possible an interim synthesis of this rapidly advancing branch of photobiology.     

Temperature-scan cryocrystallography reveals reaction intermediates in bacteriophytochrome

Light is a fundamental signal that regulates important physiological processes such as development and circadian rhythm in living organisms. Phytochromes form a major family of photoreceptors responsible for red light perception in plants, fungi and bacteria. They undergo reversible photoconversion between red-absorbing (Pr) and far-red-absorbing (Pfr) states, thereby ultimately converting a light signal into a distinct biological signal that mediates subsequent cellular responses. Several structures of microbial phytochromes have been determined in their dark-adapted Pr or Pfr states. However, the structural nature of initial photochemical events has not been characterized by crystallography. Here we report the crystal structures of three intermediates in the photoreaction of Pseudomonas aeruginosa bacteriophytochrome (PaBphP). We used cryotrapping crystallography to capture intermediates, and followed structural changes by scanning the temperature at which the photoreaction proceeded. Light-induced conformational changes in PaBphP originate in ring D of the biliverdin (BV) chromophore, and E-to-Z isomerization about the C(15) = C(16) double bond between rings C and D is the initial photochemical event. As the chromophore relaxes, the twist of the C(15) methine bridge about its two dihedral angles is reversed. Structural changes extend further to rings B and A, and to the surrounding protein regions. These data indicate that absorption of a photon by the Pfr state of PaBphP converts a light signal into a structural signal via twisting and untwisting of the methine bridges in the linear tetrapyrrole within the confined protein cavity.     

A light-sensing knot revealed by the structure of the chromophore-binding domain of phytochrome

Phytochromes are red/far-red light photoreceptors that direct photosensory responses across the bacterial, fungal and plant kingdoms. These include photosynthetic potential and pigmentation in bacteria as well as chloroplast development and photomorphogenesis in plants. Phytochromes consist of an amino-terminal region that covalently binds a single bilin chromophore, followed by a carboxy-terminal dimerization domain that often transmits the light signal through a histidine kinase relay. Here we describe the three-dimensional structure of the chromophore-binding domain of Deinococcus radiodurans phytochrome assembled with its chromophore biliverdin in the Pr ground state. Our model, refined to 2.5 A resolution, reaffirms Cys 24 as the chromophore attachment site, locates key amino acids that form a solvent-shielded bilin-binding pocket, and reveals an unusually formed deep trefoil knot that stabilizes this region. The structure provides the first three-dimensional glimpse into the photochromic behaviour of these photoreceptors and helps to explain the evolution of higher plant phytochromes from prokaryotic precursors.     

Phytochrome signalling is mediated through nucleoside diphosphate kinase 2.

Because plants are sessile, they have developed intricate strategies to adapt to changing environmental variables, including light. Their growth and development, from germination to flowering, is critically influenced by light, particularly at red (660 nm) and far-red (730 nm) wavelengths. Higher plants perceive red and far-red light by means of specific light sensors called phytochromes(A-E). However, very little is known about how light signals are transduced to elicit responses in plants. Here we report that nucleoside diphosphate kinase 2 (NDPK2) is an upstream component in the phytochrome signalling pathway in the plant Arabidopsis thaliana. In animal and human cells, NDPK acts as a tumour suppressor. We show that recombinant NDPK2 in Arabidopsis preferentially binds to the red-light-activated form of phytochrome in vitro and that this interaction increases the activity of recombinant NDPK2. Furthermore, a mutant lacking NDPK2 showed a partial defect in responses to both red and farred light, including cotyledon opening and greening. These results indicate that NDPK2 is a positive signalling component of the phytochrome-mediated light-signal-transduction pathway in Arabidopsis.     

Gating of the rapid shade-avoidance response by the circadian clock in plants

The phytochromes are a family of plant photoreceptor proteins that control several adaptive developmental strategies. For example, the phytochromes perceive far-red light (wavelengths between 700 and 800 nm) reflected or scattered from the leaves of nearby vegetation. This provides an early warning of potential shading, and triggers a series of 'shade-avoidance' responses, such as a rapid increase in elongation, by which the plant attempts to overgrow its neighbours. Other, less immediate, responses include accelerated flowering and early production of seeds. However, little is known about the molecular events that connect light perception with increased growth in shade avoidance. Here we show that the circadian clock gates this rapid shade-avoidance response. It is most apparent around dusk and is accompanied by altered expression of several genes. One of these rapidly responsive genes encodes a basic helix-loop-helix protein, PIL1, previously shown to interact with the clock protein TOC1 (ref. 4). Furthermore PIL1 and TOC1 are both required for the accelerated growth associated with the shade-avoidance response.     

Molecular basis of photoprotection and control of photosynthetic light-harvesting

In order to maximize their use of light energy in photosynthesis, plants have molecules that act as light-harvesting antennae, which collect light quanta and deliver them to the reaction centres, where energy conversion into a chemical form takes place. The functioning of the antenna responds to the extreme changes in the intensity of sunlight encountered in nature. In shade, light is efficiently harvested in photosynthesis. However, in full sunlight, much of the energy absorbed is not needed and there are vitally important switches to specific antenna states, which safely dissipate the excess energy as heat. This is essential for plant survival, because it provides protection against the potential photo-damage of the photosynthetic membrane. But whereas the features that establish high photosynthetic efficiency have been highlighted, almost nothing is known about the molecular nature of the dissipative states. Recently, the atomic structure of the major plant light-harvesting antenna protein, LHCII, has been determined by X-ray crystallography. Here we demonstrate that this is the structure of a dissipative state of LHCII. We present a spectroscopic analysis of this crystal form, and identify the specific changes in configuration of its pigment population that give LHCII the intrinsic capability to regulate energy flow. This provides a molecular basis for understanding the control of photosynthetic light-harvesting.     

拟南芥铁氧还蛋白基因缺失促进花期提前的初步研究(英文)

拟南芥的红光/远红光受体光敏色素(PHYs)参与花期调节过程,而铁氧还蛋白色素还原酶(FD-BRs)的一种——植物色素合成酶(HY2)对于光敏色素的合成是必不可少的。研究发现拟南芥铁氧还蛋白——AtFd2的基因缺失突变体(Fd2-KO突变体)在长日照与短日照培养条件下,较其野生型而言均表现出花期提前的表型,而且显示AtFd2与AtHY2在叶绿体中发生互作,并且Fd2突变体对光敏色素的反应受到抑制。推测At-Fd2基因的缺失可能通过影响光敏色素介导的相关生理功能进而对植株的花期进行调节。     

Ecology: a niche for cyanobacteria containing chlorophyll d

The cyanobacterium known as Acaryochloris marina is a unique phototroph that uses chlorophyll d as its principal light-harvesting pigment instead of chlorophyll a, the form commonly found in plants, algae and other cyanobacteria; this means that it depends on far-red light for photosynthesis. Here we demonstrate photosynthetic activity in Acaryochloris-like phototrophs that live underneath minute coral-reef invertebrates (didemnid ascidians) in a shaded niche enriched in near-infrared light. This discovery clarifies how these cyanobacteria are able to thrive as free-living organisms in their natural habitat.     

模拟干旱处理下毛竹光响应特征分析

【目的】研究毛竹叶绿素含量和光响应参数的变化,比较分析毛竹对干旱胁迫的响应特点,探讨毛竹抗旱生理与适应变化,为毛竹林的水分管理提供理论依据。【方法】以Ⅰ～Ⅲ度毛竹为试验材料,分别测定干旱和对照条件下毛竹叶片光合色素含量和光响应曲线,并计算出其最大净光合速率(P_n,max)、光饱和点(P_LSP)、光补偿点(P_LCP)、暗呼吸速率(R_d)和表观量子效率(η_AQY)等光合参数。【结果】毛竹叶片叶绿素含量在不同竹龄之间差异都不显著,而Ⅱ度竹光合速率显著高于Ⅰ度竹和Ⅲ度竹,Ⅰ度竹和Ⅲ度竹之间光合速率差异不显著; Ⅰ度竹各叶绿素含量在干旱条件下显著低于对照,而Ⅲ度竹各叶绿素含量在干旱和对照间差异不显著,Ⅱ度竹只有类胡萝卜素含量在干旱条件下显著高于对照。各竹龄毛竹叶绿素含量与光合速率变化不一致,干旱处理下Ⅰ度竹和Ⅲ度竹P_n显著高于对照,Ⅱ度竹则无显著差异。【结论】干旱处理提高了Ⅰ度竹和Ⅲ度竹光能利用效率,降低了Ⅱ度竹对光的生态适应能力。     

Functional interaction of phytochrome B and cryptochrome 2

Light is a crucial environmental signal that controls many photomorphogenic and circadian responses in plants. Perception and transduction of light is achieved by at least two principal groups of photoreceptors, phytochromes and cryptochromes. Phytochromes are red/far-red light-absorbing receptors encoded by a gene family of five members (phyA to phyE) in Arabidopsis. Cryptochrome 1 (cry1), cryptochrome 2 (cry2) and phototropin are the blue/ultraviolet-A light receptors that have been characterized in Arabidopsis. Previous studies showed that modulation of many physiological responses in plants is achieved by genetic interactions between different photoreceptors; however, little is known about the nature of these interactions and their roles in the signal transduction pathway. Here we show the genetic interaction that occurs between the Arabidopsis photoreceptors phyB and cry2 in the control of flowering time, hypocotyl elongation and circadian period by the clock. PhyB interacts directly with cry2 as observed in co-immunoprecipitation experiments with transgenic Arabidopsis plants overexpressing cry2. Using fluorescent resonance energy transfer microscopy, we show that phyB and cry2 interact in nuclear speckles that are formed in a light-dependent fashion.     

杂种金叶银杏叶片光合特性分析

【目的】 分析杂种金叶银杏子代光合特性,为进一步获得具有较强环境适应能力的金叶银杏材料提供依据。【方法】 以杂种金叶银杏(‘万年金’ב南林1号’)1年生实生苗为材料,同一母本自然授粉(父本不确定)金叶银杏1年生实生苗为对照,测定叶片光合基本参数及光响应曲线,比较二者光合能力的变化情况。【结果】 杂种金叶银杏净光合速率、气孔导度、水分利用效率均明显高于对照,表现出对季节变化更强的适应性。光响应曲线拟合结果表明:杂种金叶银杏的最大净光合速率较高,光合能力较强;同时光饱和点、内禀量子效率、光补偿点处量子效率以及表观量子效率也较高,光补偿点、暗呼吸速率也均高于对照,有机物消耗较快。相关性分析结果表明:杂种金叶银杏净光合速率与气孔导度、水分利用效率呈极显著的正相关性;气孔导度与表观量子效率、蒸腾速率呈正相关。【结论】 杂交改变了金叶银杏的光合参数,有效提高了后代对光能利用的潜能及光转化的能力,可以更好地利用全日照光,因此可以培育出对光环境适应能力更强的金叶银杏品种。     

Phototropin-related NPL1 controls chloroplast relocation induced by blue light

In photosynthetic cells, chloroplasts migrate towards illuminated sites to optimize photosynthesis and move away from excessively illuminated areas to protect the photosynthetic machinery. Although this movement of chloroplasts in response to light has been known for over a century, the photoreceptor mediating this process has not been identified. The Arabidopsis gene NPL1 (ref. 2) is a paralogue of the NPH1 gene, which encodes phototropin, a photoreceptor for phototropic bending. Here we show that NPL1 is required for chloroplast relocation induced by blue light. A loss-of-function npl1 mutant showed no chloroplast avoidance response in strong blue light, whereas the accumulation of chloroplasts in weak light was normal. These results indicate that NPL1 may function as a photoreceptor mediating chloroplast relocation.     

烤烟黄叶突变体幼苗光合特性及农艺性状分析

以烤烟黄叶突变体和通过回交转育获得的黄叶K326为实验材料,研究了烤烟黄叶突变体及黄叶K326移栽期幼苗的色素质量分数、光合特性、农艺性状、干物质积累.结果表明,烤烟黄叶突变体及黄叶K326的光合色素仅为对照的45%左右,但其光合作用效率为对照的89%左右,说明色素质量分数与光合作用效率相关性不大.同时两黄叶材料具有较高光饱点,在高光强下具有较强的光能利用率.两黄叶材料除叶厚外,其他农艺性状均优于对照,三者的干物质积累差异不显著.     

Colony development and physiological characterization of the edible blue-green alga, Nostoc sphaeroides (Nostocaceae, Cyanophyta)

The edible blue green alga, Nostoc sphaeroides Kützing, is able to form microcolonies and spherical macrocolonies. It has been used as a potent herbal medicine and dietary supplement for centuries because of its nutraceutical and pharmacological benefits. However, limited information is available on the development of the spherical macrocolonies and the environmental factors that affected their structure. This report described the morphogenesis of N. sphaeroides from single trichomes to macrocolonies. During the process the most structural features of macrocolonies on various sizes were dense maculas, rings, the compact core and the formation of liquid core; and the filaments within the macrocolonies showed different length and array depending on the sizes of macrocolonies. Meanwhile temperature and light intensity also strongly affected the internal structure of macrocolonies. As microcolonies further increased in size to form 30 mm macrocolonies, the colonies differentiated into distinct outer, middle and inner layers. The filaments of the outer layer showed higher maximum photosynthetic rates, higher light saturation point, and higher photosynthetic efficiency than those of the inner layer; whereas the filaments of the inner layer had higher content of chlorophyll a and phycobiliproteins than those of the outer layer. The results obtained in this study were important for the mass cultivation of Nostoc sphaeroides as a nutraceutical product.     

Colony development and physiological characterization of the edible blue-green alga, Nostoc sphaeroides (Nostocaceae, Cyanophyta)

The edible blue-green alga,Nostoc sphaeroides Kützing,is able to form microcolorties and spherical macrocolonies.It has been used as a potent herbal medicine and dietary supplement for centuries because of its nutraceutical and pharmacological benefits.However,lim-ited information is available on the development of the spherical macrocolonies and the environmental factors that affect their structure.This report described the morphogenesis of N.Sphaeroides from single trichomes to macrocolonies.During the process,most structural features of macrocolonies of various sizes were dense maculas,rings,the compact core and the formation of liquid core;and the filaments within the macrocolonies showed different lengths and arrays depending on the sizes of macrocolonies.Meanwhile temperature and light intensity also strongly affected the internal structure of macrocolonies.As microcolonies further increased in size to form 30 mm mac-rocolonies,the colonies differentiated into distinct outer,middle and inner layers.The filaments of the outer layer showed higher max-imum photosynthetic rates,higher light saturation point,and higher photosynthetic efficiency than those of the inner layer;whereas the filaments of the inner layer had a higher content of chlorophyll a and phycobiliproteins than those of the outer layer.The results obtained in this study were important for the mass cultivation of N.Sphaeroides as a nutraceutical product.     

Photosynthetic architecture differs in coastal and oceanic diatoms

Diatoms are a key taxon of eukaryotic phytoplankton and a major contributor to global carbon fixation. They are ubiquitous in the marine ecosystem despite marked gradients in environmental properties, such as dissolved iron concentrations, between coastal and oceanic waters. Previous studies have shown that offshore species of diatoms and other eukaryotic algae have evolved lower iron requirements to subsist in iron-poor oceanic waters, but the biochemical mechanisms responsible for their decreased iron demand are unknown. Here we show, using laboratory-cultured model species, a fundamental difference between a coastal and an oceanic diatom in their photosynthetic architecture. Specifically, the oceanic diatom had up to fivefold lower photosystem I and up to sevenfold lower cytochrome b6f complex concentrations than a coastal diatom. These changes to the photosynthetic apparatus markedly decrease the cellular iron requirements of the oceanic diatom but not its photosynthetic rates. However, oceanic diatoms might have also sacrificed their ability to acclimate to rapid fluctuations in light intensity--a characteristic of dynamic and turbid coastal waters. We suggest that diatoms, and probably other eukaryotic algal taxa, exploited this difference in the underwater light climate between oceanic and coastal waters, enabling them to decrease their iron requirements without compromising photosynthetic capacity. This adaptation probably facilitated the colonization of the open ocean by diatoms, and contributes to their persistence in this iron-impoverished environment.     

水稻STN7激酶突变体的光合生理特性分析

本实验旨在研究野生型与stn7突变体在光合性能上的差异.实验包括检测种子萌发状况、用脉冲幅度调制成像荧光计IMAG-MINI测量叶绿素含量、用SPAD-502叶绿素仪测量SPAD值以及用LI-6400XT便携式光合仪测量叶绿素荧光动力学参数并用photosynthesis软件拟合光响应曲线及相关参数.结果显示,WT和stn7植株的种子萌发情况和SPAD值几乎无差异,叶绿素荧光动力学参数显示WT的光合性能比stn7植株高,叶绿素a/b和光响应曲线的结果得出WT利用弱光的能力较强,光响应曲线结果还显示高光强下,stn7植株光合速率较高.本文研究发现STN7激酶在调节水稻的光合作用中发挥着重要作用.     

Photosystem II core phosphorylation and photosynthetic acclimation require two different protein kinases

Illumination changes elicit modifications of thylakoid proteins and reorganization of the photosynthetic machinery. This involves, in the short term, phosphorylation of photosystem II (PSII) and light-harvesting (LHCII) proteins. PSII phosphorylation is thought to be relevant for PSII turnover, whereas LHCII phosphorylation is associated with the relocation of LHCII and the redistribution of excitation energy (state transitions) between photosystems. In the long term, imbalances in energy distribution between photosystems are counteracted by adjusting photosystem stoichiometry. In the green alga Chlamydomonas and the plant Arabidopsis, state transitions require the orthologous protein kinases STT7 and STN7, respectively. Here we show that in Arabidopsis a second protein kinase, STN8, is required for the quantitative phosphorylation of PSII core proteins. However, PSII activity under high-intensity light is affected only slightly in stn8 mutants, and D1 turnover is indistinguishable from the wild type, implying that reversible protein phosphorylation is not essential for PSII repair. Acclimation to changes in light quality is defective in stn7 but not in stn8 mutants, indicating that short-term and long-term photosynthetic adaptations are coupled. Therefore the phosphorylation of LHCII, or of an unknown substrate of STN7, is also crucial for the control of photosynthetic gene expression.     

两个乌桕新品种苗木光合特性比较

[目的]探讨两个乌桕新品种'海滨绯红'、'海滨紫晶'夏季光合特性并进行比较,为乌桕新品种的栽培与推广应用提供理论依据.[方法]盆栽条件下测定2年生嫁接苗叶片光合色素含量、光合日变化、光响应曲线以及CO2响应曲线,并进行灰色关联分析.[结果]①'海滨紫晶'的叶绿素a(Chla)、叶绿素b(Chlb)、类胡萝卜素(Car)...