Unravelling proximate cues of mass flowering in the tropical forests of South‐East Asia from gene expression analyses

Elucidating the physiological mechanisms of the irregular yet concerted flowering rhythm of mass flowering tree species in the tropics requires long‐term monitoring of flowering phenology, exogenous and endogenous environmental factors, as well as identifying interactions and dependencies among these factors. To investigate the proximate factors for floral initiation of mast seeding trees in the tropics, we monitored the expression dynamics of two key flowering genes, meteorological conditions and endogenous resources over two flowering events of Shorea curtisii and Shorea leprosula in the Malay Peninsula. Comparisons of expression dynamics of genes studied indicated functional conservation of FLOWERING LOCUS T (FT) and LEAFY (LFY) in Shorea. The genes were highly expressed at least 1 month before anthesis for both species. A mathematical model considering the synergistic effect of cool temperature and drought on activation of the flowering gene was successful in predicting the observed gene expression patterns. Requirement of both cool temperature and drought for floral transition suggested by the model implies that flowering phenologies of these species are sensitive to climate change. Our molecular phenology approach in the tropics sheds light on the conserved role of flowering genes in plants inhabiting different climate zones and can be widely applied to dissect the flowering processes in other plant species.     

巴西橡胶树赤霉素信号转导途径HbRGA1结构和功能分析

DELLA蛋白是赤霉素激素信号负调控因子,具有抑制植物生长发育的作用。解析其家族成员结构与功能将有助于揭示橡胶树DELLA蛋白家族成员调控橡胶树生长发育的机制。本研究从橡胶树热研73397叶片中克隆HbRGA1的cDNA全长序列。该基因长为2136 bp,含1839 bp的ORF,编码613个氨基酸。HbRGA1蛋白序列包含DELLA和GRAS保守结构域,与杨树、木薯和橡胶树DELLA基因相似性高达82.5%。qRT-PCR分析发现HbRGA1在橡胶树叶片中表达量高,在树皮和胶乳中表达量极低。叶片中HbRGA1表达量受喷施赤霉素和脱落酸等诱导显著上调。本研究表明HbRGA1与橡胶树赤霉素等激素信号密切相关,为深入研究其在橡胶树生长发育中的结构和功能打下良好基础。     

胡杨枝芽生长特征及其展叶物候特征

以5个不同发育阶段的胡杨(Populus euphratica Oliv.)个体为研究对象,观测记录了枝芽展叶物候、枝芽生长特征和叶形变化的空间分布规律。结果表明:不同发育阶段的胡杨个体以及同一个体树冠的不同层次,其枝芽生长及其展叶物候期表现出不同的时空特征。随着树龄的增加和树冠层次的增高(由基向顶),当年新生枝条长度、枝条叶片数和叶形指数逐渐减小,但叶面积和叶片干重逐渐增大。5个不同发育阶段胡杨个体均表现出展叶物候始于树冠顶层,依次向下结束于树冠基部;展叶物候期共性表现在枝芽萌动期均在4月上旬,起始展叶期集中在4月中旬,展叶终期则在5月上旬到下旬;树龄较大的个体其枝芽萌动期、起始展叶期、展叶终期较树龄较小的个体早;其枝芽萌动期到展叶终期的时间进程较树龄较小的个体短;不同发育阶段的个体枝芽萌动期出现的时间较为离散,起始展叶期和展叶终期出现的时间较为集中。相关分析表明,出叶周期与枝条长度、枝条叶片数量和叶形指数呈极显著正相关,与叶面积和叶片干重呈显著负相关。     

Nitrogen resorption from leaves under different growth irradiance in three deciduous woody species

Resorption of nitrogen (N) from senescing leaves is an important conservation mechanism that allows plants to use the same N repeatedly. We measured the extent of N resorption in plants co-occurring in a beech forest to examine the variability of N resorption, especially in relation to growth irradiance. Measurements were done in three deciduous woody species; one adult and several juvenile trees of Fagus crenata and several adult trees of Lindera umbellata and Magnolia salicifolia. N resorption efficiency (R_EFF; percentages of leaf N that is resorbed during leaf senescence) did not differ significantly among leaves under different growth irradiances in any species we studied. R_EFF was affected by the growth stage of the tree in F. crenata with the values being consistently lower in juvenile trees than in the adult tree. N resorption proficiency (R_PROF; N concentration of dead leaves) converged to a similar value in F. crenata juvenile trees and M. salicifolia, irrespective of the presenescent leaf N concentration that was affected by growth irradiance. Again, R_PROF was lower (i.e. absolute N concentration was higher) in juvenile trees than in the adult tree in F. crenata. These results suggest that the growth irradiance does not place a great impact on the extent of N resorption, but the growth stage of the tree is influential in some species. The difference between the adult and juvenile trees may be ascribed to the size of N sink tissues, which is likely to increase with plant age.     

Genetic and reproductive consequences of forest fragmentation for populations of <Emphasis Type="Italic">Magnolia obovata</Emphasis>

In order to evaluate the consequences of forest fragmentation on populations of Magnolia obovata, we compared genetic diversity and reproductive characteristics at two nearby sites, one conserved and one fragmented. The genetic diversity between adults trees of the different sites was not significantly different. However, saplings in the conserved site showed a significantly higher genetic diversity than both adult trees in the conserved site and saplings in the fragmented sites; this was found to be the result of the larger gene flow into the conserved site. The density of the adult trees was significantly related to all of the reproductive traits analyzed (fertilization of ovules, insect attack to seeds, ovules that developed into seeds and outcrossing at the stage of seeds) at both sites. At both sites, fertilization of ovules and insect attack on seeds were positively correlated to adult tree density while outcrossing rate was negatively correlated to adult tree density. The fertilization of ovules and outcrossing were more dependent on adult tree density in the fragmented site than in the conserved site. The probability of ovules developing into outcrossed seeds showed a negative correlation with adult tree density at both sites, indicating the advantage of low density for this species and possibly implying a resilience to habitat fragmentation. A two-generation-analysis did not identify significant differences between sites in terms of the structure of the pollen pool and the number of pollen donors. Although fragmentation affected reproductive characteristics, the effect on seedling establishment and subsequent survival remains to be determined. Proposals for future studies that will assist in the development of management strategies for forests suffering fragmentation are made.     

武夷山不同龄级香果树开花物候及其生殖特性

该研究采用固定样地观测研究了不同龄级香果树的开花物候和生殖特性,分析了物候指数与生殖构件之间的相关性。结果表明：香果树单花花期一般为5~9 d,平均为6.79 d,随着树龄增大,其单花花期呈幂函数增长趋势;不同树龄香果树的始花期不同,但终花期基本一致;随着树龄的增大,始花期逐渐提前,花期持续时间延长。20~50 a树龄的香果树始花期约为8月9日,花期持续时间最短,仅为36 d。110~140 a树龄的香果树始花期约为7月15日,花期持续时间为61 d;对于单株母树来讲,香果树的花枝数,花朵总数以及果实总数随着树龄的增大均显著增加;单花枝产花量随着树龄的增加呈上升趋势,单花枝产果实数量呈先上升后下降的趋势;香果树花枝和单花枝产花量在树冠方位上的分布由大到小依次为南>东/西>北、上>中>下,但单花枝果实数量则表现出不同规律,即东>南>西>北、中>下>上。香果树具有花多果少现象,开花物候指数与生殖构件数量间的相关性分析表明,其花期持续时间与花枝数、花数和果实数均存在极显著正相关关系,始花期与生殖构件数量呈极显著负相关关系,而物候指数与香果树树龄亦存在显著相关关系。     

Constitutive expression of two apple (Malus x domestica Borkh.) homolog genes of LIKE HETEROCHROMATIN PROTEIN1 affects flowering time and whole-plant growth in transgenic Arabidopsis

Fruit trees, such as apple (Malus × domestica Borkh.), are woody perennial plants with a long juvenile phase. The biological analysis for the regulation of flowering time provides insights into the reduction of juvenile phase and the acceleration of breeding in fruit trees. In Arabidopsis, LIKE HETEROCHROMATIN PROTEIN1 (LHP1) is involved in epigenetic silencing of the target genes such as flowering genes. We isolated and characterized twin apple LHP1 homolog genes, MdLHP1a and MdLHP1b. These genes may have been generated as a result of ancient genome duplication. Although the putative MdLHP1 proteins showed lower similarity to any other known plant LHP1 homologs, a chromo domain, a chromo shadow domain, and the nuclear localization signal motifs were highly conserved among them. RT-PCR analysis showed that MdLHP1a and MdLHP1b were expressed constantly in developing shoot apices of apple trees throughout the growing season. Constitutive expression of MdLHP1a or MdLHP1b could compensate for the pleiotropic phenotype of lhp1/tfl2 mutant, suggesting that apple LHP1 homolog genes are involved in the regulation of flowering time and whole-plant growth. Based on these results, LHP1 homolog genes might have rapidly evolved among plant species, but the protein functions were conserved, at least between Arabidopsis and apple. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

续随子ElFAD2基因的序列及表达特性分析

为了研究油酸脱氢酶(FAD2)基因ElFAD2对续随子(Euphorbia lathyris L.)中不饱和脂肪酸合成的调控作用,该研究在续随子转录组数据的基础上经筛选获得ElFAD2基因序列,并对其序列及表达特性进行分析。序列分析结果显示,ElFAD2基因全长1 907 bp,ORF长1 152 bp,共编码383个氨基酸,包含有典型的脂肪酸去饱和酶结构域。续随子ElFAD2蛋白理论等电点为8.08,属于稳定蛋白,包含4个跨膜区和3个保守的组氨酸簇。基于FAD2的系统发育分析表明,续随子与同科植物乌桕(Triadica sebifera L.)的亲缘关系最近。荧光定量PCR分析发现,ElFAD2基因在不同器官中均有表达,且在花后15 d的种子中表达量最高,在叶与花后30 d及45 d种子中的表达量相当,而在根、茎、花中的表达量最低。该研究结果为深入探讨续随子ElFAD2基因的生物学功能提供了基础数据,也为解析续随子种子中脂肪酸合成的分子机制奠定了基础。     

First report of Bipolaris bicolor causing a leaf spot disease on rubber tree

The rubber tree (Hevea brasiliensis) is the only resource for commercial natural rubber production and thus has economic importance in Southeast Asia. A spot disease on the leaves of a rubber tree was first discovered in 2017 in Hainan, China. In this study, the fungal isolate MA1 from the infected tissues was determined to be a pathogen of the spot disease by satisfying Koch's postulates. The isolate MA1 was identified as Bipolaris bicolor based on the morphological characteristics and multigene phylogenetic analysis. Among fungicides, prochloraz, iprodione and pyraclostrobin significantly inhibited hyphal growth of B. bicolor under in vitro conditions. This study constitutes the first report on the association of B. bicolor with leaf spot disease of rubber trees worldwide.     

Ectopic Expression of an <Emphasis Type="Italic">FT</Emphasis> Homolog from <Emphasis Type="Italic">Citrus</Emphasis> Confers an Early Flowering Phenotype on Trifoliate Orange (<Emphasis Type="Italic">Poncirus trifoliata</Emphasis> L. Raf.)

Citrus FT (CiFT) cDNA, which promoted the transition from the vegetative to the reproductive phase in Arabidopsis thaliana, when constitutively expressed was introduced into trifoliate orange (Poncirus trifoliata L. Raf.). The transgenic plants in which CiFT was expressed constitutively showed early flowering, fruiting, and characteristic morphological changes. They started to flower as early as 12 weeks after transfer to a greenhouse, whereas wild-type plants usually have a long juvenile period of several years. Most of the transgenic flowers developed on leafy inflorescences, apparently in place of thorns; however, wild-type adult trifoliate orange usually develops solitary flowers in the axils of leaves. All of the transgenic lines accumulated CiFT mRNA in their shoots, but there were variations in the accumulation level. The transgenic lines showed variation in phenotypes, such as time to first flowering and tree shape. In F₁ progeny obtained by crossing ‘Kiyomi’ tangor (C. unshiu × sinensis) with the pollen of one transgenic line, extremely early flowering immediately after germination was observed. The transgene segregated in F₁ progeny in a Mendelian fashion, with complete co-segregation of the transgene and the early flowering phenotype. These results showed that constitutive expression of CiFT can reduce the generation time in trifoliate orange.     

Olfactory response of Stenotus rubrovittatus to rice and paddy weed,Scirpus juncoides

The olfactory response of the sorghum plant bug, Stenotus rubrovittatus (Matsumura) (Het., Miridae), to rice, Oryza sativa L., and paddy weed, Scirpus juncoides Roxb. var. ohwianus T. Koyama, was investigated with an olfactometer to clarify the mechanism of the invasion of the bugs in paddy fields. Both adult females and males were significantly attracted to panicles of rice in the flowering and full‐ripe stages. Whole plants (aboveground parts) of rice in the panicle‐formation stage, and stems and leaves of rice in the flowering stage significantly attracted only adult females. Other rice structures tested did not attract males or females. Both males and females were attracted to the flowering spikelets of S. juncoides. Although females showed no olfactory response to stems of S. juncoides in the flowering stage, males were repelled by them. Only females were attracted to whole plants of flowering S. juncoides. Whole plants of S. juncoides in the spikelet‐formation stage significantly attracted only females. The findings suggest that the invasions of S. rubrovittatus into paddy fields are caused by their olfactory responses to the volatiles emitted from rice and some paddy weeds such as S. juncoides.     

Effect of leaf age and position on light-saturated CO<Subscript>2</Subscript> assimilation rate,photosynthetic capacity,and stomatal conductance in rubber trees

Shoots of the tropical latex-producing tree Hevea brasiliensis (rubber tree) grow according to a periodic pattern, producing four to five whorls of leaves per year. All leaves in the same whorl were considered to be in the same leaf-age class, in order to assess the evolution of photosynthesis with leaf age in three clones of rubber trees, in a plantation in eastern Thailand. Light-saturated CO₂ assimilation rate (A _max) decreased more with leaf age than did photosynthetic capacity (maximal rate of carboxylation, V _cmax , and maximum rate of electron transport, J _max), which was estimated by fitting a biochemical photosynthesis model to the CO₂-response curves. Nitrogen-use efficiency (A _max/N_a, N_a is nitrogen content per leaf area) decreased also with leaf age, whereas J _max and V _cmax did not correlate with N _a. Although measurements were performed during the rainy season, the leaf gas exchange parameter that showed the best correlation with A _max was stomatal conductance (g _s). An asymptotic function was fitted to the A _max-g _s relationship, with R ² = 0.85. A _max, V _cmax, J _max and g _s varied more among different whorls in the same clone than among different clones in the same whorl. We concluded that leaf whorl was an appropriate parameter to characterize leaves for the purpose of modelling canopy photosynthesis in field-grown rubber trees, and that stomatal conductance was the most important variable explaining changes in A _max with leaf age in rubber trees.     

The coincidence of thrips and dispersed pollen in PNRSV-infected stonefruit orchards – a precondition for thrips-mediated transmission via infected pollen

Recent laboratory studies have demonstrated that Prunus necrotic ringspot virus (PNRSV) (family Bromoviridae) can be readily transmitted when thrips and virus‐bearing pollen are placed together on to test plants. For this transmission mechanism to result in stonefruit tree infection in the field, PNRSVbearing pollen must be deposited onto surfaces of stonefruit trees on which thrips also occur. In a previous paper, we demonstrated that almost all pollen in a PNRSV‐infected Japanese plum orchard in southeastern Queensland was deposited onto flowers, whereas few grains occurred on leaves and none on stems. Here, we present results of our investigation of thrips species composition, distribution and abundance on stonefruit trees in the same study area as our previous pollen deposition study. We collected a total of 2010 adult thrips from 13 orchards during the 1989, 1991 and 1992 flowering seasons of which all but 14 were in the suborder Terebrantia. Most (97.4%) terebrantian thrips were of three species, Thrips imaginis, Thrips australis and Thrips tabaci. Thrips tabaci as well as species mixtures that included T. imaginis, T. australis and T. tabaci have been shown to transmit PNRSV via infected pollen in laboratory tests. Adult thrips were frequently collected from flowers but rarely from leaves and never from stems. Large and significant differences in numbers of T. imaginis, T. australis and T. tabaci adults in flowers occurred among orchards and between seasons. No factor was conclusively related to thrips numbers but flowers of late‐flowering stonefruit varieties tended to hold more thrips than those of early‐flowering varieties. Our results indicate that the common thrips species present on stonefruit trees in the Granite Belt are also ones previously shown to transmit PNRSV via infected pollen in the laboratory and that these thrips are concentrated in tonefruit flowers where most stonefruit pollen is deposited. These results contribute to mounting circumstantial evidence that stonefruit flowers may be inoculated with PNRSV via an interaction of thrips with virus‐bearing pollen and that this transmission mechanism may be an important cause of new tree infections in the field     

Effects of flowering tree density on the mating system and gene flow in Shorea leprosula (Dipterocarpaceae) in Peninsular Malaysia

Pristine tropical rainforests in Southeast Asia have rich species diversity and are important habitats for many plant species. However, the extent of these forests has declined in recent decades and they have become fragmented due to human activities. These developments may reduce the genetic diversity of species within them and, consequently, the species’ ability to adapt to environmental changes. Our objective in the study presented here was to clarify the effect of tree density on the genetic diversity and gene flow patterns of Shorea leprosula Miq. populations in Peninsular Malaysia. For this purpose, we related genetic diversity and pollen flow parameters of seedling populations in study plots to the density of mature trees in their vicinity. The results show that gene diversity and allelic richness were not significantly correlated to the mature tree density. However, the number of rare alleles among the seedlings and the selfing rates of the mother trees were negatively correlated with the density of the adult trees. Furthermore, in a population with high mature tree density pollination distances were frequently <200 m, but in populations with low adult tree density the distances were longer. These findings suggest that the density of flowering trees affects selfing rates, gene flow and, thus, the genetic diversity of S. leprosula populations. We also found an individual S. leprosula tree with a unique reproductive system, probably apomictic, mating system.     

Spatial dispersion and sampling of citrus whitefly,Dialeurodes citri,for control decisions in a citrus orchard

1 Monitoring the citrus whitefly Dialeurodes citri (Ashmead) in citrus orchards in Israel is necessary for control decisions in IPM in the period before the establishment of natural enemies. The main damage caused by D. citri is due to sooty mould, which colonizes the sweet secretions of this pest. 2 Using Taylor’s Power Law to describe the variance–mean relationship, the dispersion of the citrus whitefly was found to be aggregated at all developmental stages. The action threshold was estimated using regression of sooty mould level on egg or larval counts. 3 Two-stage sampling plans were constructed in different pest generations, for eggs and larval stages, on the basis of the variance between leaves on a tree and the variance between trees in a plot. 4 Fixed sample size plans were prepared, with possible numbers of trees and the numbers of leaves per tree to be sampled, in order to attain a given precision level. 5 Sequential sampling plans were also drawn up, on the basis of four leaves per tree, where trees are sampled until a decision can be reached. The maximum number of trees to be sampled was set according to the corresponding fixed sample size plan. 6 Simulation of the sequential sampling plans showed them to be highly reliable in terms of the decision reached with economy in sample size for high and low infestation levels.     

Biological spectrum and other structural functional attributes of the vegetation of Kumaun Himalaya

Spectra on life form, leaf size, leaf persistence, flowering season, and shade tolerance of trees in different vegetation types occurring within the north-western catchment of the river Gola in Kumaun Himalaya are presented. The flora of Quercus leucotrichophora, Quercus lanuginosa and Quercus floribunda forests is phanerophytic, that of Pinus roxburghii and mixed forests therophytic. The grassland vegetation is characterized by the largest percentage of hemicryptophytes. The flora of the whole area, is therophytic. The biological spectrum for the entire Kumaun Himalaya is characterized as therohemigeophytic. Among the various altitudinal zones, the tropical has a preponderance of phanerophytes, the temperate and the alpine of hemicryptophytes.The observations on leaf size indicate that with the exception of Pinus roxburghii forest, in all vegetation types, the species with microphylls are greater in number. In this region, the vegetation expression is evergreen, although the tree flora has a considerable content of deciduous elements. In all the forests, the flowering period in most of the trees is vernal.On the basis of relative density, the greatest proportions of adult trees in the Pinus roxburghii and Quercus lanuginosa forests are shade intolerant, while in the mixed and Quercus leucotrichophora forests maximum trees are intermediate in shade tolerance. With the exception of the Pinus roxburghii forest, all the forests exhibit the dominance of trees which are shade tolerant at the seedling stage. On the basis of relative density, all forest types, except for Pinus roxburghii forest, have 74.5 to 100% trees with the potentiality of vegetative reproduction.Financial support from the Indian Space Research Organisation, Banglore and University Grants Commission, New Delhi is gratefully acknowledged.