基质养分对寄生植物南方菟丝子生长的影响

群落中各营养级的相互作用在群落结构形成中起了重要作用.以南方菟丝子(Cuscuta australis R.Br.)和三叶鬼针草(Biden pilosa L.)为研究对象,采用完全随机区组实验设计方法,测定并分析基质养分(不施肥与施肥)对寄生植物生长的影响,探讨寄生植物生物量与寄主生长特性、生物量和光源捕获能力的相关性.结果表明,施肥显著增加寄生植物南方菟丝子的吸器数量、缠绕圈数、相对盖度、营养器官生物量、生殖器官生物量和总生物量,但对生殖器官的生物量比无显著影响.施肥显著增加寄主植物的根、茎、叶生物量和总生物量、叶生物量比、比叶面积和叶绿素含量,但显著降低根冠比与根生物量比.南方菟丝子生物量与三叶鬼针草生物量、叶生物量比、比叶面积以及相对叶绿素含量之间均存在显著正相关,与根生物量比和根冠比存在显著负相关.研究结果表明施肥可以提高寄主植物的光资源捕获能力,将更多地生物量分配至叶等光合机构上,从而促进寄主植物(生产者)的生长,并间接促进寄生植物(初级消费者)的生长.     

Diet mixing in a parasitic plant: adaptation or constraint?

Some herbivores deliberately consume a mixed diet, either to obtain a superior mix of nutrients or to avoid consuming too much of any one toxin. Few studies have examined diet mixing in parasitic plants, which typically have very broad host ranges. We offered the parasitic plant Cuscuta indecora (dodder), a range of mixtures of two hosts (Iva frutescens and Borrichia frutescens) in the greenhouse, and observed correlations between the host community and Cuscuta infection in the field. In the greenhouse, Cuscuta performed better on mixtures with a higher relative abundance of Iva. Cuscuta selectively foraged on whichever host was more abundant (diet switching), the exact opposite of the behavior that would be expected if diet mixing was advantageous. In the field, the intensity of Cuscuta infections was decreased by the presence of non-hosts (grasses), not strongly affected by the presence of intermediate hosts, and increased by the presence of Borrichia. We conclude that Cuscuta does not obtain nutritional benefits from a broad diet, but instead is constrained by its relative lack of mobility to attack hosts of intermediate value. In general, the lack of mobility of parasitic plants compared to herbivores probably selects for broad host ranges in parasitic plants.     

Host-produced ethylene is required for marked cell expansion and endoreduplication in dodder search hyphae

The genus Cuscuta comprises stem holoparasitic plant species with wide geographic distribution. Cuscuta spp. obtain water, nutrients, proteins, and mRNA from their host plants via a parasitic organ called the haustorium. As the haustorium penetrates into the host tissue, search hyphae elongate within the host tissue and finally connect with the host’s vascular system. Invasion by Cuscuta spp. evokes various reactions within the host plant’s tissues. Here, we show that, when Arabidopsis (Arabidopsis thaliana) is invaded by Cuscuta campestris, ethylene biosynthesis by the host plant promotes elongation of the parasite’s search hyphae. The expression of genes encoding 1-aminocylclopropane-1-carboxylic acid (ACC) synthases, ACC SYNTHASE2 (AtACS2) and ACC SYNTHASE6 (AtACS6), was activated in the stem of Arabidopsis plants upon invasion by C. campestris. When the ethylene-deficient Arabidopsis acs octuple mutant was invaded by C. campestris, cell elongation and endoreduplication of the search hyphae were significantly reduced, and the inhibition of search hyphae growth was complemented by exogenous application of ACC. In contrast, in the C. campestris-infected Arabidopsis ethylene-insensitive mutant etr1-3, no growth inhibition of search hyphae was observed, indicating that ETHYLENE RESPONSE1-mediated ethylene signaling in the host plant is not essential for parasitism by C. campestris. Overall, our results suggest that C. campestris recognizes host-produced ethylene as a stimulatory signal for successful invasion.

Growth of Cuscuta campestris search hyphae is inhibited in ethylene-deficient Arabidopsis mutants, suggesting that host-derived ethylene acts as a stimulatory signal for parasitism by Cuscuta spp.     

Characterization of <Emphasis Type="Italic">PP2A</Emphasis>-<Emphasis Type="Italic">A3</Emphasis> mRNA expression and growth patterns in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> under drought stress and abscisic acid

Phosphoprotein phosphatase 2A (PP2A) plays a crucial role in cellular processes via reversible dephosphorylation of proteins. The activity of this enzyme depends on its subunits. There is little information about mRNA expression of each subunit and the relationship between these gene expressions and the growth patterns under stress conditions and hormones. Here, mRNA expression of subunit A3 of PP2A and its relationship with growth patterns under different levels of drought stress and abscisic acid (ABA) concentration were analyzed in Arabidopsis thaliana. The mRNA expression profiles showed different levels of the up- and down-regulation of PP2AA3 in roots and shoots of A. thaliana under drought conditions and ABA treatments. The results demonstrated that the regulation of PP2AA3 expression under the mentioned conditions could indirectly modulate growth patterns such that seedlings grown under severe drought stress and those grown under 4 µM ABA had the maximum number of lateral roots and the shortest primary roots. In contrast, the minimum number of lateral roots and the longest primary roots were observed under mild drought stress and 0.5 µM ABA. Differences in PP2AA3 mRNA expression showed that mechanisms involved in the regulation of this gene under drought conditions would probably be different from those that regulate the PP2AA3 expression under ABA. Co-expression of PP2AA3 with each of PIN1-4,7 (PP2A activity targets) depends on the organ type and different levels of drought stress and ABA concentration. Furthermore, fluctuations in the PP2AA3 expression proved that this gene cannot be suitable as a reference gene although PP2AA3 is widely used as a reference gene.     

Evidence for abscisic acid biosynthesis in Cuscuta reflexa, a parasitic plant lacking neoxanthin

Abscisic acid (ABA) is a plant hormone found in all higher plants; it plays an important role in seed dormancy, embryo development, and adaptation to environmental stresses, most notably drought. The regulatory step in ABA synthesis is the cleavage reaction of a 9-cis-epoxy-carotenoid catalyzed by the 9-cis-epoxy-carotenoid dioxygenases (NCEDs). The parasitic angiosperm Cuscuta reflexa lacks neoxanthin, one of the common precursors of ABA in all higher plants. Thus, is C. reflexa capable of synthesizing ABA, or does it acquire ABA from its host plants? Stem tips of C. reflexa were cultured in vitro and found to accumulate ABA in the absence of host plants. This demonstrates that this parasitic plant is capable of synthesizing ABA. Dehydration of detached stem tips caused a big rise in ABA content. During dehydration, ¹⁸O was incorporated into ABA from ¹⁸O₂, indicating that ABA was synthesized de novo in C. reflexa. Two NCED genes, CrNCED1 and CrNCED2, were cloned from C. reflexa. Expression of CrNCEDs was up-regulated significantly by dehydration. In vitro enzyme assays with recombinant CrNCED1 protein showed that the protein is able to cleave both 9-cis-violaxanthin and 9′-cis-neoxanthin to give xanthoxin. Thus, despite the absence of neoxanthin in C. reflexa, the biochemical activity of CrNCED1 is similar to that of NCEDs from other higher plants. These results provide evidence for conservation of the ABA biosynthesis pathway among members of the plant kingdom.Abscisic acid (ABA) is found in all higher plants and algae and is also produced by some fungi (Oritani and Kiyota, 2003; Schwartz and Zeevaart, 2004; Nambara and Marion-Poll, 2005). In higher plants, ABA is involved in seed dormancy, embryo development, and adaptation to various abiotic stresses. ABA is a sesquiterpenoid (C₁₅). In some fungi, there is a direct pathway from isopentenyl pyrophosphate (C₅) via farnesyl pyrophosphate (C₁₅; Oritani and Kiyota, 2003). Higher plants synthesize ABA via the C₄₀ indirect pathway. A C₄₀ carotenoid is oxidatively cleaved to form a C₂₅ byproduct and the C₁₅ precursor of ABA, xanthoxin. Biochemical and molecular evidence has shown that the cleavage reaction is the rate-limiting step in the ABA biosynthetic pathway.Although the pathway of ABA biosynthesis in higher plants has been well established, there are still a few unresolved questions. One is the endogenous substrate of the cleavage reaction. The biochemical evidence has indicated that the C₄₀ substrate for production of biologically active ABA is an epoxy-carotenoid in the 9-cis configuration in order for biologically active ABA to be produced. In higher plants, the major 9-cis-epoxy-carotenoids are 9′-cis-neoxanthin and 9-cis-violaxanthin. Because 9′-cis-neoxanthin is the most abundant 9-cis-epoxy-carotenoid in higher plants, it has been speculated that 9′-cis-neoxanthin is the main endogenous substrate of ABA. However, in vitro enzyme assays with recombinant 9-cis-epoxy-carotenoid dioxygenase (NCED) proteins from several plant species have shown that NCEDs are capable of cleaving both 9′-cis-neoxanthin and 9-cis-violaxanthin (Schwartz et al., 1997; Qin and Zeevaart, 1999; Chernys and Zeevaart, 2000), with a higher activity when 9-cis-violaxanthin is used as substrate (Schwartz et al., 2003b). This raised the question whether one or both are the endogenous substrates of ABA biosynthesis.Cuscuta reflexa is of interest in addressing this question, because it lacks 9′-cis-neoxanthin, one of the major 9-cis-epoxy-carotenoids found in green plants. A study by Bungard et al. (1999) investigated the xanthophyll-carotenoid complement of the main light-harvesting complex involved in light capture of photosynthesis. They found that in most higher plants, the complex is highly conserved and includes, in addition to chlorophyll, the carotenoids neoxanthin, violaxanthin, and lutein. In C. reflexa, neoxanthin is replaced by another xanthophyll, lutein-5,6-epoxide. Therefore, C. reflexa lacks the step that converts violaxanthin to neoxanthin. There are two xanthophyll cycles in C. reflexa: under excess irradiance violaxanthin and lutein-5,6-epoxide are converted to zeaxanthin and lutein, respectively, by de-epoxidation; the cycle is reversed under low irradiance (Bungard et al., 1999).C. reflexa is parasitic on the above-ground parts of other plants. The leafless stems coil around the host stems and petioles. C. reflexa contains only a small amount of chlorophyll and relies on obtaining photosynthetic assimilates from the phloem of its host. It forms specialized structures, called haustoria, which are used to acquire nutrients from its host plant (Hibberd et al., 1998).The lack of 9′-cis-neoxanthin in C. reflexa may result in an inability of the plant to produce ABA, and the parasite may acquire it from its host, thereby eliminating the need to synthesize ABA on its own (Bungard et al., 1999). The objective of this study was to determine whether C. reflexa is able to synthesize ABA or acquires it from its host. Increased ABA in C. reflexa grown independently of a host would indicate that it is capable of ABA synthesis. We also searched for the presence of an NCED gene in the Cuscuta genome. Presence of an NCED gene would be evidence for the 9-cis-epoxy-carotenoid cleavage reaction, the regulatory step for ABA synthesis. Our results show that despite the lack of 9′-cis-neoxanthin, C. reflexa is capable of ABA synthesis in a manner similar to that in other higher plants.     

The parasitic mechanism of the holostemparasitic plant Cuscuta

Cuscuta is a stem holoparasitic plant without leaves or roots, which develops a haustorium and sucks nutrients from host plants. The genus Cuscuta comprises about 200 species, many of which can cause severe problems for certain crops. The parasitic process in Cuscuta begins in finding and attaching to a host plant and then developing a haustorium. The process does not always require any chemical signal, but does require a light signal. Finding a host involves detecting the lower red light:far-red light ratio near a potential host plant by phytochrome. A contact signal is also necessary for haustorium induction. Apparently, cytokinin increase is downstream of the light and contact signal and is critical for haustorium induction. This pathway, however, appears to be slightly different from a standard pathway. The direct connection between Cuscuta and its host involves both the xylem and phloem, and mRNA and proteins can translocate. Several features indicate that Cuscuta is a useful model plant for parasite plant research as well as plant–plant interaction research. These include the simple anatomical structure and seedling development, no chemical requirement for haustorium induction, and the wide range of host plants.     

Responses of antioxidant defense system of <Emphasis Type="Italic">Helianthus annuus</Emphasis> to abscisic acid treatment under drought and waterlogging

The effect of abscisic acid (ABA) treatment on growth pigments and antioxidant defense system were investigated in seedlings of Helianthus annuus (cvs. Nantio F1 and Özdemirbey) subjected to drought and waterlogging stress. In addition, seedlings were sprayed with 10 M ABA three times every other day. Relative growth rate (RGR) was significantly reduced in both genotypes under drought stress, however, this growth inhibition was less in ABA-treated plants. Total chlorophyll content increased by drought stress in both genotypes. Ascorbate was not influenced by drought, while α-tocopherol increased in cv. Nantio F1. Ascorbate and α-tocopherol increased with drought stress in cv. Özdemirbey. ABA treatment decreased ascorbate and β-carotene contents while it increased α-tocopherol and xanthophylls contents under drought stress. The activity of superoxide dismutase (SOD) in both genotypes increased under drought stress-ABA combinations. Catalase (CAT) activity decreased under drought stress and drought-ABA combinations while it increased under waterlogging stress. Glutathione reductase (GR) activity decreased under drought stress but recovered with ABA treatment. The results suggested that ABA treatments have different effects on the components of antioxidant defense system in H. annuus genotypes and ABA may contribute drought-induced oxidative stress tolerance but not effects under waterlogging stress.     

Plant Responses to Drought Stress and Exogenous ABA Application are Modulated Differently by Mycorrhization in Tomato and an ABA-deficient Mutant (<Emphasis Type="Italic">Sitiens</Emphasis>)

The aims of the present study are to find out whether the effects of arbuscular mycorrhizal (AM) symbiosis on plant resistance to water deficit are mediated by the endogenous abscisic acid (ABA) content of the host plant and whether the exogenous ABA application modifies such effects. The ABA-deficient tomato mutant sitiens and its near-isogenic wild-type parental line were used. Plant development, physiology, and expression of plant genes expected to be modulated by AM symbiosis, drought, and ABA were studied. Results showed that only wild-type tomato plants responded positively to mycorrhizal inoculation, while AM symbiosis was not observed to have any effect on plant development in sitiens plants grown under well-watered conditions. The application of ABA to sitiens plants enhanced plant growth both under well-watered and drought stress conditions. In respect to sitiens plants subjected to drought stress, the addition of ABA had a cumulative effect in relation to that of inoculation with G. intraradices. Most of the genes analyzed in this study showed different regulation patterns in wild-type and sitiens plants, suggesting that their gene expression is modulated by the plant ABA phenotype. In the same way, the colonization of roots with the AM fungus G. intraradices differently regulated the expression of these genes in wild-type and in sitiens plants, which could explain the distinctive effect of the symbiosis on each plant ABA phenotype. This also suggests that the effects of the AM symbiosis on plant responses and resistance to water deficit are mediated by the plant ABA phenotype.     

Cytokinin-producing, plant growth-promoting rhizobacteria that confer resistance to drought stress in Platycladus orientalis container seedlings

One of the proposed mechanisms through which plant growth-promoting rhizobacteria (PGPR) enhance plant growth is the production of plant growth regulators, especially cytokinin. However, little information is available regarding cytokinin-producing PGPR inoculation on growth and water stress consistence of forest container seedlings under drought condition. This study determined the effects of Bacillus subtilis on hormone concentration, drought resistance, and plant growth under water-stressed conditions. Although no significant difference was observed under well-watered conditions, leaves of inoculated Platycladus orientalis (oriental thuja) seedlings under drought stress had higher relative water content and leaf water potential compared with those of noninoculated ones. Regardless of water supply levels, the root exudates, namely sugars, amino acids and organic acids, significantly increased because of B. subtilis inoculation. Water stress reduced shoot cytokinins by 39.14 %. However, inoculation decreased this deficit to only 10.22 %. The elevated levels of cytokinins in P. orientalis shoot were associated with higher concentration of abscisic acid (ABA). Stomatal conductance was significantly increased by B. subtilis inoculation in well-watered seedlings. However, the promoting effect of cytokinins on stomatal conductance was hampered, possibly by the combined action of elevated cytokinins and ABA. B. subtilis inoculation increased the shoot dry weight of well-watered and drought seedlings by 34.85 and 19.23 %, as well as the root by 15.445 and 13.99 %, respectively. Consequently, the root/shoot ratio significantly decreased, indicative of the greater benefits of PGPR on shoot growth than root. Thus, inoculation of cytokinin-producing PGPR in container seedlings can alleviate the drought stress and interfere with the suppression of shoot growth, showing a real potential to perform as a drought stress inhibitor in arid environments.     

Antioxidant response and Lea genes expression under exogenous ABA and water deficit stress in wheat cultivars contrasting in drought tolerance

Two wheat (Triticum aestivum) cultivars, C306 (drought tolerant) and PBW343 (drought susceptible) were compared for their response to exogenous ABA, water stress (WS) and combined (ABA plus WS) during their seedlings growth. Their responses were studied in the form of seedlings growth, antioxidant potential of roots and shoots and expression levels of LEA genes in shoots. ABA treatment led to increase in levels of ascorbate, ascorbate to dehydroascorbate ratio, antioxidant enzymes and decreases in levels of dehydroascorbate, malondialdehyde (MDA). Decrease in biomass, ascorbate contents, ascorbate to dehydroascorbate ratios and antioxidant enzymes was more in PBW343 than in C306 under WS. Dehydroascorbate and MDA levels were higher in PBW343 than in C306 under WS. ABA plus WS improved some of these features from their levels under WS in PBW343. Proline contents were not increased significantly under ABA in both cultivars. Out of ten LEA genes studied, six LEA genes were induced more under WS than under ABA in C306 but equally induced in PBW343. Four LEA genes were induced earlier in PBW343 but later in C306. Wdhn13 was induced more under ABA than under WS in C306 while it was non-responsive to both stresses in PBW343.     

空心莲子草响应南方菟丝子寄生的生长-防御权衡

为探讨全寄生植物南方菟丝子(Cuscuta australis)防治入侵植物空心莲子草(Alternanthera philoxeroides)的可行性,以二者野外天然生长的种群为研究对象,分析南方菟丝子寄生对空心莲子草生长及防御的影响,阐明空心莲子草在受到寄生胁迫时如何权衡自身生长与防御的关系,进而发展出一套应对南方菟丝子寄生的生长-防御策略。结果显示:(1)南方菟丝子寄生显著改变空心莲子草茎的形态,茎直径和平均节间长均增加,茎直径变化极显著(P0.01);(2)南方菟丝子寄生显著减少空心莲子草叶片数,但同时显著增加后者茎的分枝数,而茎上的节是潜在的无性繁殖体,故有利于空心莲子草的克隆繁殖;此外,南方菟丝子寄生显著降低了空心莲子草的根、茎、叶生物量和总生物量,抑制空心莲子草的生长;(3)南方菟丝子寄生显著增加空心莲子草茎的单宁、总酚、三萜皂苷含量,增强其防御能力;(4)南方菟丝子寄生的空心莲子草的生物量与茎部木质素、三萜皂苷、单宁和总酚含量均呈现显著负相关性(P0.01),对照组则不存在相关性;且寄生组较对照组相比,生物量的相对百分比显著低于对照组(P0.01),而用于防御的次生代谢产物总含量的相对百分比显著高于对照组(P0.01)。以上结果表明,受到南方菟丝子寄生胁迫后,空心莲子草改变自身的生长-防御策略,减少营养生长投入而将更多的资源投向克隆繁殖,同时增强对"防御"物质的投入,增强其防御能力,以利于后代生存和繁衍。     

自然干旱胁迫下长春花叶片和根部的激素和可溶性糖代谢变化

植物在离开生长环境较短时间内（1~6 h）会导致缓慢的表面水分散失,引起自然的干旱胁迫。本文以耐旱植物长春花（Catharanthus roseus）为材料,研究其在离土干旱胁迫中的脱落酸（ABA）及可溶性糖含量变化。结果表明,长春花根部ABA含量在正常条件下低于叶片中的含量,干旱胁迫促进了ABA在根部的积累,6 h时增加至最高值。蔗糖酸性转化酶活性可能受到ABA的诱导在胁迫6 h时最高,比对照高出30%左右。长春花叶片中总可溶性糖含量在对照条件下非常稳定,但在干旱胁迫过程中,其随着时间的延长呈现线性增加的趋势（r²=0.964）,蔗糖和已糖含量在胁迫过程中也呈增加的趋势,可能发挥着渗透调控节功能。     

Metabolite changes with induction of Cuscuta haustorium and translocation from host plants

Cuscuta is a stem holoparasitic plant without leaves or roots, parasitizing various types of host plants and causing major problems for certain crops. Cuscuta is known as a generalist and, thus, must have unique parasite strategies to cope with different host plants. For elucidating metabolic responses and mechanisms of parasitization, metabolomic approaches using GC/MS were applied. We compared five stages of Cuscuta japonica: early stage seedlings, with far red light (FR) cue, with contact signal, haustorium induced seedlings by both signals and adult plant parasites on host plants. Sugars, amino acids, organic acids, nucleic acids, and polyols were identified from the polar phase fraction. The apical part contained metabolite profiles different from the haustorium induced part or the basal part. Amino acid and some organic acids were up-regulated for haustorium induction but decreased after parasitization. After attachment to different host plants, metabolite profiles of Cuscuta japonica changed dramatically due to the absorption of specific host plant metabolites such as pinitol. Cuscuta seedlings attached to pinitol rich host plants contained more pinitol and showed different profiles from those attached to plants having less or lacking pinitol.     

A chromosome-scale Gastrodia elata genome and large-scale comparative genomic analysis indicate convergent evolution by gene loss in mycoheterotrophic and parasitic plants

Mycoheterotrophic and parasitic plants are heterotrophic and parasitize on fungi and plants, respectively, to obtain nutrients. Large-scale comparative genomics analysis has not been conducted in mycoheterotrophic or parasitic plants or between these two groups of parasites. We assembled a chromosome-level genome of the fully mycoheterotrophic plant Gastrodia elata (Orchidaceae) and performed comparative genomic analyses on the genomes of G. elata and four orchids (initial mycoheterotrophs), three parasitic plants (Cuscuta australis, Striga asiatica, and Sapria himalayana), and 36 autotrophs from various angiosperm lineages. It was found that while in the hemiparasite S. asiatica and initial mycoheterotrophic orchids, approximately 4–5% of the conserved orthogroups were lost, the fully heterotrophic G. elata and C. australis both lost approximately 10% of the conserved orthogroups, indicating that increased heterotrophy is positively associated with gene loss. Importantly, many genes that are essential for autotrophs, including those involved in photosynthesis, the circadian clock, flowering time regulation, immunity, nutrient uptake, and root and leaf development, were convergently lost in both G. elata and C. australis. The high-quality genome of G. elata will facilitate future studies on the physiology, ecology, and evolution of mycoheterotrophic plants, and our findings highlight the critical role of gene loss in the evolution of plants with heterotrophic lifestyles.     

The effect of nutrients on seedling growth of native and introduced Phragmites australis

Differing responses to abiotic stresses and increased nutrient availability may play a role in the invasion and spread of introduced Phragmites australis Cav. (Trin.) ex. Steud. and the decline of native P.a. americanus Saltonstall, P.M. Peterson & Soreng in North America. We present results from an outdoor experiment where native and introduced P. australis seedlings were grown under two nutrient treatments. Both subspecies responded positively to increased nutrients but introduced plants clearly outperformed natives, growing taller, producing more stems, and had three to four times higher biomass. The biomass of introduced P. australis growing in low nutrients was similar to that of the native in high nutrients. Aboveground:belowground biomass ratios were nearly 1.25 for both native and introduced plants across treatments and reflect the high investment P. australis seedlings place on shoot production in their first year of growth. Our results also demonstrate that introduced P. australis can have explosive growth over a single growing season, even when established from seed. This implies that management of young, newly established populations may be prudent where introduced P. australis is considered undesirable, irregardless of whether eutrophication is an issue.