Photoperiodism dynamics during the domestication and improvement of soybean

Soybean(Glycine max) is a facultative short-day plant with a sensitive photoperiod perception and reaction system, which allows it to adjust its physiological state and gene regulatory networks to seasonal and diurnal changes in environmental conditions. In the past few decades, soybean cultivation has spread from East Asia to areas throughout the world. Biologists and breeders must now confront the challenge of understanding the molecular mechanism of soybean photoperiodism and improving agronomic traits to enable this important crop to adapt to geographical and environmental changes. In this review, we summarize the genetic regulatory network underlying photoperiodic responses in soybean. Genomic and genetic studies have revealed that the circadian clock, in conjunction with the light perception pathways, regulates photoperiodic flowering. Here, we provide an annotated list of 844 candidate flowering genes in soybean, with their putative biological functions. Many photoperiod-related genes have been intensively selected during domestication and crop improvement. Finally, we describe recent progress in engineering photoperiod-responsive genes for improving agronomic traits to enhance geographic adaptation in soybean, as well as future prospects for research on soybean photoperiodic responses.

Photoperiod Mediates the Effects of Temperature and Light Intensity on the Proliferation of Ulva prolifera

In order to study the complex effects of photoperiod,temperature,and light intensity on the spore maturation and release number of Ulva prolifera,we cultured thalli segment(2–3 mm)under three different photoperiods(L:D=12:12,14:10 and 10:14),temperature(15℃(LT),25℃(MT)and 30℃(HT))and light intensity(100,200 and 400μmol m^(−2)s^(−1),noted as LL,ML and HL,respectively)conditions.Then the maturation time,spore release number and chlorophyll fluorescence were analyzed.The results suggested that:1)The spore maturation time was accelerated by higher temperature or higher light intensity from 62 h to 36 h,and changes in day length accelerated the spore maturation to a certain extent as compared with 12:12 light/dark cycle;2)Higher light intensity significantly decreased the chlorophyll fluorescence(Fv′/Fm′,NPQ,rETRmax andα)of the mature reproductive segment under 30℃with 12:12 light/dark cycle.But when in the other photoperiods(10:14 and 14:10 conditions),the inhibitory effects of high light intensity were alleviated significantly;3)The optimum condition for the spore maturation and release was 12:12 light/dark cycle,25℃,400μmol m^(−2)s^(−1),with both shorter and longer photoperiod reducing the spore release number;4)Higher light intensity significantly increased the spore release number under 25℃,but these effects were alleviated by 30℃treatment.This study is the first attempt to elucidate the coincidence effects of photoperiod,temperature and light intensity on the reproduction of Ulva,which would help to reveal the mechanism of the rapid proliferation of green tide.

Flowering-time regulation by the circadian clock:From Arabidopsis to crops

Precise timing of flowering in plants is critical for their growth and reproductive processes.One factor controlling flowering time is the cycle of light and darkness within a day,known as the photoperiod.Plants are classified into long-day,short-day,and day-neutral plants based on light requirements for floral initiation.Although the molecular mechanisms that govern this differentiation remain incompletely understood,studies have consistently shown that the circadian clock plays a central role in regulating photoperiod response across diverse plant species.However,there is a scarcity of reviews describing the regulatory network linking the circadian clock with photoperiodic flowering.This review summarizes that regulatory network,focusing on the distinct roles of clock genes in long-day and short-day plants.We also discuss the strategies of clock gene mutations contributing to geographic variation in longday and short-day crops.

Molecular mechanism of flowering time regulation in Brassica rapa:similarities and differences with Arabidopsis

Properly regulated flowering time is pivotal for successful plant reproduction.The floral transition from vegetative growth to reproductive growth is regulated by a complex gene regulatory network that integrates environmental signals and internal conditions to ensure that flowering takes place under favorable conditions.Brassica rapa is a diploid Cruciferae species that includes several varieties that are cultivated as vegetable or oil crops.Flowering time is one of the most important agricultural traits of B.rapa crops because of its influence on yield and quality.The transition to flowering in B.rapa is regulated by several environmental and developmental cues,which are perceived by several signaling pathways,including the vernalization pathway,the autonomous pathway,the circadian clock,the thermosensory pathway,and gibberellin(GA)signaling.These signals are integrated to control the expression of floral integrators BrFTs and BrSOC1s to regulate flowering.In this review,we summarized current research advances on the molecular mechanisms that govern flowering time regulation in B.rapa and compare this to what is known in Arabidopsis.

Effects of heat stress and long photoperiod on the prostate of rats

Objective:To examine light and heat effects on the morphological,histological,and micrometric structure of the prostate of rats.Methods:Thirty adult male rats were divided into three groups.The control group was kept under 20℃-22℃ and an artificial 12 h/12 h day/night cycle;the temperature group was under normal light and at(42±1)℃ heat for 4 to 5 h daily,and the light group was exposed to 8 h/16 h day/night cycle with 20℃-22℃.Rats were weighed five times(at the beginning of the study and every seven days).Five milliliters(mL)of their peripheral blood were taken.The tissue staining was performed using the hematoxylin-eosin(H&E)stain and periodic acid-Schiff(PAS).In the following,tissue and cellular reactions to the PAS were examined.Results:Folds were located entirely on the surface of the anterior lobe and periphery of the other lobes.The secretory units in the anterior lobe were more than the lateral lobe.A strong reaction of the secretory cells to the PAS was observed.Testosterone serum levels of the light group also significantly increased compared to the control group(P<0.05).The most histometric changes of the lobes were established in the lateral lobes.Heat stress resulted in a significant decrease in testosterone levels and transformed prostate tissue.The epithelium and parenchyma to scaffold ratio in the temperature group decreased.Conclusions:Maximum and minimum changes in the ventral lobe happened under the ascent of temperature and light,respectively.The ventral lobe in the study of prostatic hyperplasia should be more considered.

FLOWERING LOCUS T(FT)in Photosensitive Type Chrysanthemum Accelerates Flowering in Arabidopsis

FLOWERING LOCUS T(FT)is an important factor for integrating flowering signals through the photoperiod pathway,which significantly promotes flowering.Most chrysanthemum varieties are short-day plants,sensitive to the photoperiod,which limits their ornamental and annual production supply.In this study,we aim to analyze the nutrients and flowering-related genes of chrysanthemums with different photoperiod types and to clone and verify the function of the flowering-related gene CmFT.We found that the formation of floral buds requires the accumulation of starch while consuming soluble sugars and the expression patterns of flowering-related genes GIGANTEA(GI),CONSTANS(CO),and FT in C.morifolium‘Zilian’and C.morifolium‘Zihongtuogui’had a certain synchronization during floral buds differentiation according to our quantitative validation,and the expression levels of CmGI,CmCO and CmFT in C.morifolium‘Zihongtuogui’were higher than those in C.morifolium.‘Zilian’in the later stage of differentiation.CmFT was cloned from photosensitive chrysanthemums-C.morifolium‘Zihongtuogui’and polypeptide alignment and phylogenetic analysis showed that CmFT was clustered in FT-like subfamily.In further functional verification,we obtained two Arabidopsis transgenic lines.Our results showed that CmFT transgenic ft mutant lines can significantly accelerate flowering in Arabidopsis.Thus,we can initially confirm that CmFT plays an important role in promoting flowering,which may be the key reason for the photosensitivity of C.morifolium‘Zihongtuogui’.Overall,the results of this study are of great importance in revealing the flowering mechanism of different photoperiod types of chrysanthemums.

Common Organic Amendment (Rice Straw) Can Reduce Salinity Effects on Bean (Phaseolus vulgaris) Growth with or without Photoperiod Manipulation

Soil salinity is a major limiting factor for crop production in coastal areas of Bangladesh. Cheap and sustainable management of soil salinity is hence most sought out topics in agricultural research. Conceptualizing that idea in mind, a pot experiment was conducted in the Department of Soil, Water & Environment, University of Dhaka in order to analyze if common organic amendments (rice straw, saw dust) coupled with reduce photoperiod can mitigate salinity effect on the growth of bean (Phaseolus vulgaris). The experiment was set up following completely randomized design (CRD) with nine treatments and three replications containing Tc (Control), T1 (Ambient photoperiod + 110 mM Salinity treatment + Rice straw), T2 (Reduced photoperiod + 110 mM Salinity treatment + Rice straw), T3 (Ambient photoperiod + 220 mM Salinity treatment + Rice straw), T4 (Reduced photoperiod + 220 mM Salinity treatment + Rice straw), T5 (Ambient photoperiod + 110 mM Salinity treatment + Saw dust), T6 (Reduced photoperiod + 110 mM Salinity treatment + Saw dust), T7 (Ambient photoperiod + 220 mM Salinity treatment + Saw dust) and T8 (Reduced photoperiod + 220 mM Salinity treatment + Saw dust). Organic amendments were used separately at the rate of 12 ton/ha. The highest plant height (98.67 cm), root length (12.5 cm), pod number (10.33), leaf area (13.99 cm2), fresh weight (680 kg/ha), dry weight (316.67 kg/ha) were recorded with the treatment T1 while the second-best treatment was treatment T2 (with highest harvest index 0.040) and these results were statistically significant (p < 0.001). In post-harvest soil, pH, EC, OC, OM;available N, P, K, S;total Ca, Mg, Zn, Mn were increased significantly in treatment T1. The overall results illustrated that the best growth and yield performances were achieved in the treatment T1 and T2.

Effect of Shortening Day Length on Immune Function in Siberian Hamsters

In order to understand the effect of gradual changes in photoperiod on immune function, adult female Siberian hamsters (Phodopus sungorus) were randomly divided into the control group (12L:12D, Con, n = 11) and the shortening day length group (SD, n = 11), in which day length was reduced from 12:12 h to 8:16 h light-dark cycle at the pace of half an hour every week. Meanwhile the winter immunoenhancement hypothesis, which holds that animals’ immune function would be enhanced in winter or winter-like conditions, was tested. Gradual shortening day length had no effect on body mass and body composition including wet carcass mass, the subcutaneous, retroperitoneal, mesenteric and total body fat masses in Siberian hamsters. The masses of liver and small intestine with contents were higher in the SD group than in the Con group, however other organ masses such as brain, heart, kidney and so on did not differ between the two groups. Phytohemagglutinin (PHA) response after 24 h of PHA injection was enhanced by the shortening photoperiod, which supported the winter immunoenhancement hypothesis. The masses of spleen and thymus, white blood cells, bacteria killing capacity indicative of innate immunity were not affected, which did not support this hypothesis. In summary, gradually decrease in day length increased cellular immunity, but had no effect on other immunological parameters in Siberian hamsters.

Immunohistochemical Analysis of IAA in Anthers of the Photoperiod-sensitive Genic Male-sterile Rice

The immunohistochemical localization of IAA and the comparison of their relative levels were carried out for the first time in the anthers of Nongken 58S and its wild type Nongken 58 (Oryza sativa subsp. japonica) after long_day and short_day treatments. The distribution of free_IAA in anthers and its dynamic variation could be reflected by this method. The results showed that the IAA level in the anthers of Nongken 58S after long_day treatment was much lower than that in short_day_treated Nongken 58S and those in wild type Nongken 58 in five stages from pistil and stamen primordia formation to late uninucleate stage. The possible reasons for IAA deficiency in Nongken 58S_LD anthers and its relationship with fertility alteration were also discussed.

Comparative Studies on the Changes of Microtubule Distribution and Reorganization During the Meiotic Stages of Development in Normal (IR36) and a Temperature/photoperiod Sensitive Male Sterile Line (Peiai 64S) of Rice ( Oryza sativa )

Changes in the pattern of organization of microtubules in the meiotic stages of development of pollen (i.e. from pre-meiotic interphase to more or less metaphase I) of a normal (IR36) and a temperature/photoperiod sensitive male sterile line (Peiai 64S) of rice were studied using immunofluorescence confocal microscopy. In IR36, from pre-meiotic interphase to metaphase I, the pattern of microtubule distribution in the meiocytes underwent a series of changes. Some new organizational patterns of microtubules (that have not been described before) were observed during microsporogenesis, including the existence of a broad band of perinuclear microtubules at the diakinesis stage of development. The pattern of microtubule distribution in the meiocytes of the male sterile line, Peiai 64S, was quite different front that seen in IR36. In Peiai 64S, the microtubules showed abnormal patterns of distribution from pre-meiotic interphase to metaphase I. For example the broad band of perinuclear microtubules seen at diakinesis in IR36 was much disorganized and loosened in Peiai 64S. The spindles formed were also very abnormal and different from the normal spindle. The appearance of abnormal microtubule distribution in the early stages of microsporogenesis may contribute to the malformation and ultimate abortion of pollen in Peiai 64S.

Preliminary Studies on the Effect of Photoperiod and Temperature on Male Fertility of a Cytoplasmic Male Sterile Line in Wheat

[Objective] The study aimed to reveal the effect of photoperiod and temperature on male fertility of cytoplasmic male sterile line Vtai911289a in wheat and discuss the mechanism of male fertility alteration. [ Method ] The sowing-date tests and designed conditions were conducted during 2003 -2005. [ Result] Fertility of Vtai911289a, could alter under specific photoperiod and temperature conditions. Temperature is one of the main factors influencing male fertility of the male sterile lines. Vtai911289a, showed stable sterility under the condition of the mean of daily temperature at fertility sensitive stage lower than 19℃ and presented partial fertility when the mean of daily temperature at fertility sensitive stage lower than 20 - 22℃. Photoperiod to some extent affects the male fertility of Vtai911289a, long-day condition is helpful for the male fertility of the sterile line. [ Conclusion] The application of photoperiod temperature-sensitive cytoplasmic male sterile line in production has a higher safety than that of temperature sensitive sterile line.

Photoperiod Control of Apical Bud and Leaf Senescence in Pumpkin ( Cucurbita pepo ) Strain 185

Short_day (SD) induced the plant senescence in pumpkin ( Cucurbita pepo Linn.) strain 185. Structural assay, gene expression and a series of biochemical analyses were performed to analyze the senescence mechanism in pumpkin strain 185 exposed to SD. Two aspects of important changes initiated in SD exposure contributed to the senescence process. SD functionally led to the initiation of the apical transformation from vegetative to reproductive growth, and then programmed cell death (PCD) in the apical meristem, causing the loss of vigorous growth activity. Moreover, SD treatment resulted in the formation of a great number of dying cells in mesophyll tissue later in the development compared with the phenotype of plants under long_day (LD) conditions. During the senescence process, high expression of nuclease is an important molecular event. These results indicate that the initiation of senescence process in pumpkin stain 185 plants is closely related to the death of cells in apical meristem and mesophyll.

Reproductive polyphenism and its advantages in aphids:Switching between sexual and asexual reproduction

Reproductive polyphenism,which allows one genotype to produce sexual and asexual morphs,is an extreme case of phenotypic plasticity and is commonly observed in aphids.Aphids are typical species that switch these reproductive modes,and the pathway orientation is triggered by the environmental conditions(mainly photoperiod and temperature).The typical arm ual life of aphids in eludes a successi on of parthenogenetic gen erations duri ng the spri ng and summer and a single sexual generation in autumn.In this review,we describe how the environmental cues orientate the reproductive mode of aphids from photoperiodic perception to endocrine regulation,and how juvenile hormones may act on the target cells(oocytes)to initiate the gametogenesis and embryogenesis in sexual and asexual reproduction.We also discuss the paradox of sex,especially the advantages of sexual reproduction in aphids.With the recent development of genomic resources in aphids,many potential genes involved in the reproductive polyphenism will enter the public's awareness.In particular,we describe a novel RNAi method in aphids,which may provide a molecular technique for determining the developmental fate and multiple reproductive strategies.

Advances in Melatonin and Its Functions in Plants

Abstract Melatonin （N-acetyi-5-methoxytryptamine） is a well-known animal hormone, which is synthesized and secreted by pineal gland and takes part in the regulation of circadian rhythm in animals. At present it has been observed that melatonin is widely existed in higher plants while there have no enough studies on functions of melatonin in plants. Researches have already indicated that the possible functions of melatonin in plants include regulating photoperiod, participating in growth regulation, clearing active oxygen, and promoting activity of antioxidase. General reviews upon functions of melatonin in plants are made upon experiments in recent years. We fo-cus on the demonstrated and predicted biological functions of melatonin in plants to bring researchers up to date on this field. The weakness in present studies and the main research directions are also pointed out.

The development of the Rook's(Corvus frugilegus)colony in the face of shifting spring phenology

Many birds species breed in colonies.One of the species in the Corvidae family nesting in groups is the Rook(Corvus frugilegus).The construction of the nest is costly for birds and needs high energy expenditure.Therefore,birds should optimize the time of nest building in relation to environmental,especially weather,conditions.Furthermore,birds should adapt their breeding phenology,including the date of starting the construction of nests for climate change.We observed the dynamics of increasing numbers of nests in a colony of Rooks in relation to air temperature,wind,rainfall,snowfall and day-length(indirect indicator of photoperiodism).Observations were carried out during three breeding seasons(2015-2017)in a large rookery in a big city in north-eastern Poland.The increase in the number of nests was correlated only with day length and no effect of climatic factors on the number of nests in the colony was observed.Number of nests varied significantly between successive years of research.In the case of the Rook(predictable breeders),the photoperiod seems to be the most important factor influencing breeding behavior,especially nest construction.In the last three decades,increasing temperature and time of spring phases have advanced and have influenced changes in the phenology of the availability of food resources.Simultaneously,a rapid decline in the numbers of breeding pairs of the Rook in Poland has been observed,especially in the last three decades.Conservative attachment to the length of the day,which is the main factor responsible for the development of the breeding colony,makes the Rook appear to be a species with low plasticity in the face of rapid climate changes.Thus,we suggest that may indirectly influence the decrease in the breeding population of the Rook.

Effects of photoperiod on daily activity rhythm of juvenile sea cucumber, Apostichopus japonicus (Selenka)

In nature, Apostichopus japonicus exhibits a behavioral characteristic of emerging at night and sheltering during the day. Hence, it was commonly believed that longer and darker light conditions are better for a population of A. japonicus. In aquaculture situations therefore, animals have commonly been cultured without lighting, especially during juvenile-hatching culture. However, how the length of darkness each day affects .4.japonicus and what the mechanism is of behavioral response of A. japonicus to different photoperiods remain unclear, and are specifically addressed in this study. We applied nine photoperiod treatments （light：dark （LD） cycles at 0：24, 3：21, 6：18, 9：15, 12：12, 15：9, 18：6, 21：3, and 24：0 under 500Ix） to 60 individuals （29.73±0.23 g） per treatment （with 3 replicates）. The enviroramental conditions were set as： water temperature = 16.9±0.3℃, dissolved oxygen〉6.0 rag/L, ammonia〈0.3 mg/L, pH=7.8-8.2, and salinity=30-32. Under experimental conditions of continuous darkness as well as continuous light, cyclic nocturnal activity patterns of A. japonicus （viz. the animals emerged and fed at night and sheltered during the day） were observed. However, they spent more and less time, respectively, moving and feeding under continuous darkness and continuous light, than those under a natural light cycle. Under photoperiods with 6-12 h of light, the animals showed one sheltering behavior transition and two emerging behavior transitions. These behavior transitions appeared to be governed by some internal physiological factors and induced by daily light variation. The behavior of the animals was significantly affected by different photoperiods, and the distribution rate （DR） of the animals emerging at ＂daytime＂ increased with lengthening ＂light time＂ （except for the LD 3：21 cycle）. We also found that there was no significant difference in mean DR per day under LD cycles of 6：18, 9：15, 15：9, and 12：12. Mean DRs per day were significantly lower for these photoperiods, than for those under LD cycles of 0：24 and 3：21, and significantly higher than for those under LD cycles of 18：6, 21：3 and 24：0. These results imply that 6-15 h light per day were the similar photoperiods to natural sunlight. From the viewpoint of activity and feeding behaviors, these might be the optimal photoperiods for rearing A. japonicus.

Growth and phosphate uptake kinetics of Microcystis aeruginosa under various environmental conditions

Growth and uptake of exogenous phosphate by Microcystis aeruginosa in batch culture under different temperature, photoperiod, and turbulence were studied by the method of phosphate isotope tracer. Relatively high temperature, long photoperiod and strong turbulence increased the cell density of M. aeruginosa in these batch cultures. The initial rapid uptake of phosphate by M. aeruginosa was independent of the temperature, photoperiod, and turbulence. Similarly, maximum exogenous phosphate uptake was not related to these environmental factors. However, elevated temperature and turbulence shortened the time, required to obtain maximum P accumulation. The growth of M. aeruginosa could alleviate the phosphorous leakage. Total amounts of exogenous phosphate uptake to M. aeruginosa and the phosphorus leakage of M. aeruginosa were significantly influenced by the growth state of M. aeruginosa closely correlated with the environmental factors. The maximum volume of exogenous phosphate uptake to M. aeruginosa was 46% of added exogenous phosphate in water with 16 hours of photoperiod. Thus, total amounts of exogenous phosphate uptake to M. aeruginosa were more strongly affected by the photoperiod length than temperature and turbulence.

Thoughts and Practice on Some Problems about Research and Application of Two-Line Hybrid Rice

The main problems about research and application of two-line hybrid rice were reviewed, including the confusing nomenclature and male sterile lines classification, the unclear characteristics of photoperiod and temperature responses and the unsuitable site selection for male sterile line and hybrid dce seed production. In order to efficiently and accurately use dual-purpose genic male sterile lines, four types, including PTGMS （photo-thermo-sensitive genic male sterile rice）, TGMS （thermo-sensitive genic male sterile rice）, reverse PTGMS and reverse TGMS, were proposed. A new idea for explaining the mechanism of sterility in dual-purpose hybrid rice was proposed. The transition from sterile to fertile was involved in the cooperative regulation of major-effect sterile genes and photoperiod and/or temperature sensitive ones. The minor-effect genes with accumulative effect on sterility were important factors that affected the critical temperature of sterility transfer. In order to make better use of dual-purpose lines, the characterization of responses to photoperiod and temperature of PTGMS should be made and the identification method for the characterization of photoperiod and temperature responses of PTGMS should also be put forward. The optimal ecological site for seed production could be determined according to the historical climate data and the requirements for the meteorological conditions during the different periods of seed production.

CmFTL2 is involved in the photoperiod- and sucrose-mediated control of flowering time in chrysanthemum

The chrysanthemum genome harbors three FT-like genes:CmFTL1 and CmFTL3 are thought to act as regulators of floral induction under long-day(LD)and short-day(SD)conditions,respectively,whereas the function of CmFTL2 is currently unclear.The objective of the present research was to explore the function of CmFTL2 in the determination of flowering time of the photo-insensitive chrysanthemum cultivar‘Floral Yuuka’,both in response to variation in the photoperiod and to the exogenous provision of sucrose.Spraying leaves of‘Floral Yuuka’plants with 50 mM sucrose accelerated flowering and increased the level of CmFTL2 transcription in the leaf more strongly than either CmFTL1 or FTL3 under both long and SD conditions.Transcription profiling indicated that all three CmFTL genes were upregulated during floral induction.The relationship of the CmFTL2 sequence with that of other members of the PEBP family suggested that its product contributes to the florigen rather than to the anti-florigen complex.The heterologous expression of CmFTL2 in the Arabidopsis thaliana ft-10 mutant rescued the mutant phenotype,showing that CmFTL2 could compensate for the absence of FT.These results suggest that CmFTL2 acts as a regulator of floral transition and responds to both the photoperiod and sucrose.

CCT family genes in cereal crops: A current overview

Control of flowering time is crucial for reproductive success of cereal crops, and has a significant impact on grain yield as well as adaptation to diverse environmental conditions.Plants integrate signals from both environmental cues and endogenous regulatory pathways to fine-tune flowering time. The CCT domain originally described to a 43-amino acid sequence at the C-terminus of three Arabidopsis proteins, namely CONSTANS(CO),CO-LIKE, and TIMING OF CAB1(TOC1). The CCT domain-containing genes(CCT genes),which encode transcription co-factors, are the major genetic determinants that modulate flowering time, and this in turn enables plants to effectively expand their territory to take advantage of favorable habitats. Moreover, certain CCT genes have pleiotropic effects on morphological traits and confer resistance/tolerance to biotic/abiotic stresses. CCT genes can be classified into three families, namely COL(CONSTANS-like), PRR(Pseudo-response regulator), and CMF(CCT motif family),based on their non-CCT domains. During domestication, natural and artificial selection resulted in reduced nucleotide diversity of CCT genes in modern cultivated cereals than their wild types. Here, we review the features and functions of CCT genes in cereal crops and propose future research to focus on CCT genes and their utilization in crop breeding.