Progress in parasitic plant biology: host selection and nutrient transfer

Host range varies widely among species of parasitic plants. Parasitic plants realize host selection through induction by chemical molecular signals, including germination stimulants and haustoria-inducing factors (HIFs). Research on parasitic plant biology has provided information on germination, haustorium induction, invasion, and haustorial structures and functions. To date, some molecular mechanisms have been suggested to explain how germination stimulants work, involving a chemical change caused by addition of a nucleophilic protein receptor, and direct or indirect stimulation of ethylene generation. Haustorium initiation is induced by HIFs that are generated by HIF-releasing enzymes from the parasite or triggered by redox cycling between electrochemical states of the inducers. Haustorium attachment is non-specific, however, the attachment to a host is facilitated by mucilaginous substances produced by haustorial hairs. Following the attachment, the intrusive cells of parasites penetrate host cells or push their way through the host epidermis and cortex between host cells, and some types of cell wall-degrading enzymes may assist in the penetration process. After the establishment of host-parasite associations, parasitic plants develop special morphological structures (haustoria) and physiological characteristics, such as high transpiration rates, high leaf conductance, and low water potentials in hemiparasites, for nutrient transfer and resource acquisition from their hosts. Therefore, they negatively affect the growth and development and even cause death of their hosts.     

Haustoria in action: Case studies of nitrogen acquisition by woody xylem-tapping hemiparasites from their hosts

Summary This review discusses studies conducted by the author and his colleagues on mistletoes and root hemiparasites native to Western Australia. Morphological characteristics of haustoria are described and their anatomical features are discussed in relation to uptake, transfer, and metabolism of xylem-borne nitrogenous solutes derived from a host. Experimental approaches used include comparisons of xylem sap composition of parasite and host(s), solute pool analyses and enzymatic properties of haustoria, host xylem feeding of¹⁵N-labelled solutes to follow the fate of label in haustoria and body of the parasite, and studies using species-specific nonprotein amino acids to validate successful uptake from hosts or occasional backflow of xylem-borne solutes to a host. Field studies on promiscuous root hemiparasites assess frequencies of exploitation of different hosts.¹⁵N natural abundance assays of host and parasite dry matter demonstrate marked preference bySantalum acuminatum for N₂-fixing as opposed to nonfixing hosts. The ability ofOlax phyllanthi to continue to exploit deep-rooted hosts ranks of importance when xylem water potentials of other hosts go out of sucking range during periods of water stress. Comparisons of xylem sap composition of parasites feeding on different hosts indicate remarkable versatility by haustoria in uptake and utilization of the different major nitrogenous solutes received from these hosts. Solute pools in parasites partly reflect metabolic transformations accentuated by haustoria while also indicating direct throughput from xylem of a host. The review concludes by showing how empirically based modelling techniques can be used to estimate proportional gains of N by parasites from single hosts and repercussions on host growth which accompany such exploitation.Dedicated to Professor Brian E. S. Gunning on the occasion of his 65th birthday     

Heterotrophic gain of carbon from hosts by the xylem-tapping root hemiparasite Olax phyllanthi (Olacaceae)

Heterotrophic gains of carbon from various host species by the root hemiparasitic shrub Olax phyllanthi (Labill) R.Br. were assessed using techniques based on carbon isotope discrimination (¹³C) on C3 and C4 hosts and C:N ratios of xylem sap and dry matter of host and parasite. Heterotrophic benefits (H) to Olax based on ¹³C values were 30% and 19% from two nonnative C4 hosts (Portulaca oleracea and Amaranthus caudatus respectively) compared with 13% and 15% from these hosts when computed on the basis of C:N ratios of host xylem sap and C and N increments of Olax dry matter. Nitrate was the source of N available to pot cultures of the above species and estimates based on C:N ratios assumed that all N accumulated by Olax had come from nitrate absorbed by the host. Equivalent estimates of H for Olax, grown in nitrogen-free pot culture with the native N₂-fixing host Acacia littorea as its sole source of N, indicated 63% and 51% dependence on host carbon when assessed in terms of xylem sap composition of host parasite respectively. Comparisons of xylem sap solutes of Olax and a range of partner hosts indicated marked selectivity in haustorial uptake and transfer of nitrate, amino compounds, organic acids and sugars. Possible implications of variations between hosts in absolute levels of C and inorganic and organic forms of N in xylem are discussed in relation to evidence of much better growth performance of Olax on Acacia littorea and other N₂-fixing legumes than on non-fixers.     

Leaf carbon isotope and mineral composition in subtropical plants along an irradiance cline

Summary Leaf carbon isotope ratios and leaf mineral composition (Ca, K, Mg, Mn, N, and P) were measured on the dominant species along an irradiance cline in a subtropical monsoon forest of southern China. This irradiance cline resulted from disturbance caused by fuel-harvesting. Leaf carbon isotope ratios increased from undisturbed to disturbed sites for all species, indicating that leaf intercellular CO₂ concentrations decreased and leaf water use efficiencies increased along this cline. Nitrogen and magnesium levels were lower in leaves of species on the disturbed sites, but there were no clear patterns for calcium, potassium, phosphorus or manganese.C.I.W.D.P.B. Publication no 932     

Water use efficiency and carbon isotope composition of plants in a cold desert environment

Summary The effects of the availabilities of water and nitrogen on water use efficiency (WUE) of plants were investigated in a sagebrush steppe. The four species studied wereArtemisia tridentata (shrub),Ceratoides lanata (suffrutescent shrub),Elymus lanceolatus (rhizomatous grass), andElymus elymoides (tussock grass). Water and nitrogen levels were manipulated in a two-by-two factorial design resulting in four treatments: control (no additions), added water, added nitrogen, and added water and nitrogen. One instantaneous and two long-term indicators of WUE were used to testa priori predictions of the ranking of WUE among treatments. The short-term indicator was the instantaneous ratio of assimilation to transpiration (A/E). The long-term measures were 1) the slope of the relationship between conductance to water vapor and maximum assimilation and 2) the carbon isotope composition (¹³C) of plant material. Additional water decreased WUE, whereas additional nitrogen increased WUE. For both A/E and ¹³C, the mean for added nitrogen alone was significantly greater than the mean for added water alone, and means for the control and added water and nitrogen fell in between. This ranking of WUE supported the hypothesis that both water and nitrogen limit plant gas exchange in this semiarid environment. The short- and long-term indicators were in agreement, providing evidence in support of theoretical models concerning the water cost of carbon assimilation.     

多年放牧对不同类型草原植被及土壤碳同位素的影响

放牧是草原牧区常见的人类活动,多年放牧对草原植被及土壤的碳过程产生较大的影响.本研究采集不同类型草原多年放牧前后植被及土壤样品,对室内碳同位素进行分析,研究了不同草原生态系统Δ¹³C(碳同位素分馏值)差异及其影响因素.结果表明: 放牧强度对植被Δ¹³C值的影响显著,0～5 cm表层土壤Δ¹³C值在放牧前后变化显著,而对深层土壤(>5 cm)影响不显著.多年放牧后大部分植被Δ¹³C值显著升高,高海拔地区升高的幅度较大.可见,放牧行为对不同草地生态系统类型、不同土壤深度以及不同海拔生态系统碳过程产生的影响差异显著.针对不同类型的草原,放牧应采取多样化的管理方式.     

Interactions between hemiparasitic plants and their hosts: The importance of organic carbon transfer

Hemiparasitic plants display a unique strategy of resource acquisition combining parasitism of other species and own photosynthetic activity. Despite the active photoassimilation and green habit, they acquire substantial amount of carbon from their hosts. The organic carbon transfer has a crucial influence on the nature of the interaction between hemiparasites and their hosts which can oscillate between parasitism and competition for light. In this minireview, we summarize methodical approaches and results of various studies dealing with carbon budget of hemiparasites and the ecological implications of carbon heterotrophy in hemiparasites.Key words: haustorium, heterotrophy, parasitic plant, mistletoe, Rhinanthus, Striga, δ¹³CHemiparasitic plants withdraw resources from the vascular system of their hosts through a specialized transfer organ called haustorium.¹ Hemiparasites attack the host''s xylem, in contrast to the holoparasites that infect both phloem and xylem, and as a consequence, hemiparasitic plants have access to water and mineral nutrients but little carbon.¹ Due to their reduced or non-existing root networks, hemiparasitic plants acquire virtually all mineral nutrients and water from the host while organic carbon is provided, at least in part, by their own photosynthetic activity.^2,3 This is in contrast to holoparasitic plants which rely on the host for the supply of both organic and inorganic nutrients. The location of the attachment to the host and the degree of host dependency represent the most important characters defining the three basic functional types within hemiparasitic plants. Root hemiparasites attack host roots but their above-ground appearance is usually not substantially different from that of a non-parasitic plant. This group can be further divided in two—facultative and obligate hemiparasites consisting of plants that are able (at least sometimes) or unable to complete their life cycle without an attachment to the host respectively. Stem hemiparasites are attached to the host stem (usually trunk or branches) and are all obligate parasites, unable to survive without a host.Hemiparasitic plants have an ambiguous relationship with their hosts which, on the one hand, represent exclusive sources of inorganic nutrients but on the other hand, the co-occurrence of these host plants in the hemiparasite vicinity imposes competition for light. The nature and intensity of this competitive relationship varies across different groups and species of hemiparasites. The ability of hemiparasites to acquire organic carbon (largely in the form of xylem-mobile organic and amino acids) is certainly the key factor affecting this interaction since hemiparasites that are capable of efficient organic carbon abstraction should be minimally affected by shading from their host. The fact that hemiparasites can exhibit substantial carbon heterotrophy is now supported by a large number of studies, although a traditional point of view on hemiparasites that highlights the importance of inorganic resources (mainly nitrogen) acquisition is still prevailing. Therefore, we decided to summarize available information on hemiparasite heterotrophy, outline techniques for assessing the proportion of heterotrophy and estimating the overall carbon budget, and discuss possible implications of this phenomenon on hemiparasite ecology.     

Telomere dynamics in parasitic great spotted cuckoos and their magpie hosts

Although little is known on the impact of environment on telomere length dynamics, it has been suggested to be affected by stress, lifestyle and/or life‐history strategies of animals. We here compared telomere dynamics in erythrocytes of hatchlings and fledglings of the brood parasite great spotted cuckoos (Clamator glandarius) and of magpies (Pica pica), their main host in Europe. In magpie chicks, telomere length decreased from hatching to fledging, whereas no significant change in telomere length of great spotted cuckoo chicks was found. Moreover, we found interspecific differences in the association between laying date and telomere shortening. Interspecific differences in telomere shortening were interpreted as a consequence of differences in lifestyle and life‐history characteristics of magpies and great spotted cuckoos. In comparison with magpies, cuckoos experience reduced sibling competition and higher access to resources and, consequently, lower stressful environmental conditions during the nestling phase. These characteristics also explain the associations between telomere attrition and environmental conditions (i.e. laying date) for magpies and the absence of association for great spotted cuckoos. These results therefore fit expectations on telomere dynamics derived from interspecific differences in lifestyle and life history of brood parasites and their bird hosts.     

双向标记培养植物测定大气二氧化碳稳定碳同位素组成

基于植物能够利用体内的碳酸酐酶来催化碳酸氢根离子生成二氧化碳和水作为底物进行光合作用的特性,采用两种δ13CPDB值差值大于10‰的碳酸氢钠分别作为外源碳酸氢根离子的碳同位素标记物,通过室内双向水培诸葛菜和芥菜型油菜实验,分别向水培处理液里添加已知δ13CPDB值的碳酸氢钠并培养24 h,利用同位素比值质谱(IRMS)技术,测定并计算了两个时间、两种环境下的大气二氧化碳稳定碳同位素日平均组成。结果表明:在环境1(不同浓度的Na HCO3处理液)下所得到的δCa值与添加到处理液中的碳酸氢根离子的浓度有关;在环境2(不同浓度的PEG处理液)下所得到的δCa值与添加到处理液中的PEG的浓度无关;两种环境下所测得的大气二氧化碳稳定碳同位素日平均组成δCa值与实验中培养的植物种类无关,而与添加到培养液中碳酸氢根离子的浓度及植物的生长速率有关。数据重现性好,结果准确可靠,可以高精度的测定不同待测环境下大气二氧化碳稳定碳同位素比值,其可为以后监测不同时间、不同地点的大气二氧化碳碳同位素组成及来源提供非常有效的方法和信息。     

Stable isotope composition of water in desert plants

A survey of the stable isotope content of tissue waters of plants from the Negev desert was conducted. Large differences were observed in the extent of enrichment of the heavy isotopes in leaf water relative to local precipitation among different plants. This is apparently caused by the species-dependent stratagems adopted by the plants to cope with water stress, primarily by differences in the depth of water uptake in the soil and through the timing of stomatal openings during the daily cycle. Salt stressed plants showed extreme variability in the isotopic composition of leaf–water. The results show that plants with adaptation to arid conditions can avoid the transpiration regime, which would lead to the strong isotopic enrichment in their leaf water expected under arid conditions. This has implications for the use of stable isotopes in plants as indicators of either plant ecophysiology or paleoclimate. Responsible Editor: Hans Lambers. G. Goodfriend is deceased.     

C3植物稳定碳同位素组成与盐分的关系

植物在盐生环境中δ13C值的改变可能包含两个成分:一个是盐分对CO2的扩散、传递或光合速率的影响而引起的δ13C值的改变;另一个是光合途径的转换引起的δ13C值的变化,δ13C值的大小与诱导发生CAM或C4代谢的程度有关.植物组织的δ13C值随盐度的变化趋势除了与植物本身固有的耐盐性有关以外,盐度和胁迫时间是影响植物δ13C的重要因素.根据盐生条件下同位素分馏特点可知,盐生植物与非盐生植物的δ13C随盐度的变化趋势有所不同.对非盐生植物而言,在低盐度和短期的盐处理下,随盐度的增加和胁迫时间的延长植物的δ13C值增大,这个阶段限制光合作用的主要因素是气孔导度;但是如果盐度过低,δ13C变化很小,则难以表现出应有的相关性;随着胁迫的加强,当限制光合作用的非气孔因素成为主导因素时,由于光合作用受到强烈抑制(光合结构遭到破坏),δ13C将随之降低.对盐生植物而言,其δ13C与最适盐度有关.最适盐度下,植物的δ13C低于其它盐度条件下的δ13C值.盐生条件下,有些C3植物可能发生光合途径的转换,无论诱导发生的是C4代谢还是CAM代谢,δ13C值均趋于增大.但是,一般情况下,盐处理诱导的光合途径的改变对植物组织整体的δ13C的影响很小.在密闭环境中或郁闭林地,植物和土壤呼吸释放的CO2再次参与光合作用,也会改变植物的δ13C值.为了更加全面地考察植物δ13C与盐度的关系,需要设置较大的盐度范围和进行长期的胁迫处理,才能够获得相对充分的数据,才有利于全面分析植物δ13C值与耐盐性的关系.     

云南哀牢山两种常见半寄生植物的生态化学计量特征及其与寄主的关系

为深入探究半寄生植物与其寄主间的养分关系, 在云南哀牢山徐家坝地区选取两种常见半寄生植物椆树桑寄生(Loranthus delavayi)和柳叶钝果寄生(Taxillus delavayi), 研究其寄主枝条-吸器-寄生枝条-寄生叶片间的碳(C)、氮(N)、磷(P)生态化学计量特征关系。结果表明: 1)两种半寄生植物的寄主枝条-吸器-寄生枝条-寄生叶片这一连续体的C、N、P生态化学计量特征变化趋势并不完全相同, 具有物种自身的特性。2)同一半寄生植物的寄主枝条具有相似的C、N、P生态化学计量特征, 寄主物种对半寄生植物的生态化学计量特征没有显著影响。3)寄主枝条的C、N、P生态化学计量特征间具有紧密的相关关系, 吸器弱于寄主枝条, 寄生枝条弱于吸器, 寄生叶片的N、P含量相关关系不显著。4)寄主枝条与寄生叶片间的C含量存在极显著负相关关系。5)吸器与寄主枝条间的C、N、P生态化学计量特征存在紧密的相关关系, 但在生态化学计量特征的数值上吸器与寄生枝条更为接近。吸器作为连接寄主与寄生植物的关键部位, 其与寄主枝条极为显著的相关关系体现了寄主枝条养分对寄生植物的重要性, 而吸器在元素含量、计量比的数值以及相互关系上与寄生枝条更为相似, 则体现了吸器作为寄生植物器官具有与寄生枝条相似的生理功能。通过对寄主枝条-吸器-寄生枝条-寄生叶片C、N、P生态化学计量特征的分析, 为深入研究半寄生植物的养分利用策略与生态适应性提供了重要的基础资料。     

寄生植物对寄主的选择和影响

寄生植物对寄主的选择性与寄主营养物质的含量、合成的次生物质和硬度有关．多寄主存在的条件下,有利于寄生植物的生长发育．通过改变寄主光合产物的流向和影响寄主叶片的气孔调节,寄生植物对寄主生理和形态产生显著的影响．寄生植物的存在使植物群落的生物量、组成多样性和动态发生改变．大量研究显示．寄生植物对寄主的选择和影响与植食性动物对取食植物的选择和影响有很多相似之处．气候变化对寄生植物与寄主关系影响的研究还刚刚起步,寄生植物对寄主选择和影响的研究对有害寄生植物的防除和有益寄生植物的利用有重要的价值,应该重视和加强．     

Water relations of the root hemiparasite Olax phyllanthi (Labill) R.Br. (Olacaceae) and its multiple hosts

Summary Water relations of the root hemiparasite Olax phyllanthi were compared with those of its major species of hosts in natural habitat in coastal heath near Denmark, SW Australia. Leaf water potentials of Olax during winter were 0.4 to 1.4 MPa lower (more negative) than those of all (29) non parasitic host species examined. During the dry summer months (January to March), shallow-rooted hosts developed water potentials up to 3 MPa lower than those of Olax, and were accordingly rated as no longer accessible as a source of water to the hemiparasite. By contrast, deep-rooted hosts, with access to the water table, showed water potentials less negative than Olax, and haustorial contacts retained with these apparently enabled continued extraction of water and nutrients throughout the summer. Three other species of root hemiparasites parasitized by Olax, but not themselves parasitizing Olax, showed leaf water potentials throughout the year very close to, and mostly slightly more negative than those of Olax. Nocturnal measurements of leaf water potential in winter (July and August) in soil at field capacity (water potential –0.006 MPa) showed maintenance of a 0.5–0.8 MPa potential difference between Olax and a range of common host species. By dawn most hosts had equilibrated with the water potential of the soil, whereas both exposed and bagged Olax plants recorded potentials of –0.8 MPa. Daytime rates of transpiration and photosynthesis of Olax were less than those of a range of common hosts, but water use efficiencies were not consistently different between hemiparasite and hosts. This was reflected in almost identical mean values for carbon isotope ratio (¹³C/¹²C) between Olax (mean value –27.0) and thirteen frequently exploited hosts ( value –27.1). The results are discussed in relation to published information on other angiosperm hemiparasites.     

Physical and chemical basis of carbon isotope fractionation in plants

Naturally-occurring variations in the abundances of the stable isotopes of carbon and other elements can be used to understand the dynamics of natural processes in chemistry, biochemistry, biology, medicine, ecology and other fields. The use of carbon-13 isotopic abundances as an indicator of photosynthetic function in plants has become common. The purpose of this article is to describe the physical and chemical processes that contribute to the abundances of carbon-13 in plant materials, and to provide a framework for understanding how those processes control the isotopic contents of natural materials.     

Some peculiarities of the relationships between parasitic copepods and their invertebrate hosts

According to the rule of academican E. N. Pavlovskiy, any organism of host is an environment of inhabit for a parasite (Pavlovskiy, 1934). It was analysed, which "ecological niche" or microbiotop (= microhabitat) is occupied by this or that species of symbiotic (parasitic) copepods in organisms of different groups invertebrate-hosts. The assumption lying in a basis of the given analysis means that each group of hosts may give to cohabitants only certain variants of microbiotopes independently on the general morphological structure and life mode of hosts. Five types of microbiotops offered by various groups of hosts for symbiotic copepods are designated (Ta[symbol: see text] 2). 1. The body surface of benthic invertebrates as a microbiotope is characterized by conditions being little different (concerning any kind of physical and chemical influences on copepods) from those in external environment on any other substrate. Apparently a trophical dependence plays a determining role in this case. There are certain directions in a development of adaptations, which are characteristic in some extent for all water ectoparasitic crustaceans and have one functional task--to help to an ectoparasite to keep itself on a surface of host body. In the first, the maxillules and maxillipeds significantly are developed, they get a form of large claws, with which the dopepods are strongly attached on a surface of host body and have an opportunity to move on it without a danger to be washed off. In the second, the form of the body undergoes a dorso-ventral expression and expansion of prosome, forms a cephalic shield allowing to the symbiont to press itself tightly to the host body surface and to avoid the loss of host (tab. 2). In occasions, some ectoparasites stimulate the formation of galls in skin tissues of the host, that also provides the parasite with constant conditions, without any threat to lose the host. However, this phenomenon has not a wide distribution and is observed in some groups of crustacean and echinoderm hosts. 2. The narrow tubular cavities in the organism of host either they are a part of external environment (as in channel system of spongia) or a part of internal environment of organism (as channels of blood system or thin parts of a digestive system) have always rigidly limited sizes and form. Characteristics of all parasites occupying this microbiotopes are the strong transformations. They are expressed by the reduction of legs or any other appendages (frequently in a significant degree), loss of segmentation to some extent and in eruciform (or vermiform) form of a body (tab. 2). This microbiotope is occupied by an ectoparasite in one case only (Spongicola uncifer from channel system of spongia) and by endoparasites in all other cases. 3. Large cavities connected with external environment. The formations of various genesis, such as mantle cavity of molluscs, gill cavity and marsupium of crustaceans, bursal cavity of ophiuroids and branchial cavity of ascidians, concern this type of microbiotopes. All of them are characterized by the relative difference from the external environment and rather large volume (in comparison to sizes of copepods), that provides the parasites with a sufficient protection from factors of the external environment and constant source of food such as elements of host body or food's particles brought by the water flow. Morphological changes in inhabitants of the microbiotope have two directions. They practically are absent in the overwhelming majority copepods, living in the mantle of cavity of the lamellibranches. On the other hand, the inhabitants of gill cavity and marsupium of crustaceans, bursal cavities of ophiuroids and branchial cavity of ascidians are characterized by the presence of strong transformations. Usually there are expressed in a loss of segmentation to some extent, reduction of appendages and swelling of body, as in species of the genus Sphaeronella (tab. 2). Changes are also observed in the life cycle: the tendency to reduce stages of development (development of nauplii stage, which takes place under the ovarial cover). In this case the copepodid stages hatch from the ova. 4. The internal cavity of organism of host. This type of microbiotopes in different groups of the hosts is represented in a various degree. We recognise it in a coelome of polychaetes, lacunar system of molluscs, mixocoel of crustaceans, coelome of echinoderms and cavity of body in ascidians. Two basic evolutionary directions are observed in copepods occupying this microbiotope. In the first case, the parasite is not exposed to transformations and keeps the initial plan of structure as in ancestral free-living forms. In the second case the parasites are exposed to strong transformations, they either live directly in cavity's liquid, or are surrounded by a cyst (as in Cucumaricolidae). 5. Microbiotope of the last type is most specific. The simultaneous existence in two environments--external environment (environment of the second order) and internal environment (environment of the first order) leads to the complete loss of ancestral type in a structure and level of organisation. At the same time both morphological and functional division of the parasite body into two parts produces a new formation--the ectosome and endosome. In this case we deals with the phenomenon of mesoparasitism.     

Correlations between carbon isotope ratio and microhabitat in desert plants

Summary Water is usually considered to be the key limiting factor for growth of desert plants, yet there is little information available of the water-use efficiency of species within a desert community. Leaf carbon isotope ratios, an indicator of long-term intercellular carbon dioxide concentrations and thus of water-use efficiency in C₃ plants, were measured on species occurring within a Sonoran Desert community, consisting of wash, transition and slope microhabitats. Along a soil moisture gradient from the relatively wetter wash to the relatively drier slope, leaf carbon isotope ratios increased in all species, indicating that water-use efficiency increased as soil water availability decreased. Leaf carbon isotope ratios of long-lived perennials were substantially more positive than in short-lived perennials, even though plants were growing adjacent to each other. Leaf carbon isotope ratio and leaf duration (evergreen versus deciduous) were not correlated with each other. The results are discussed in terms of how the efficiency of water use may affect community structure and composition.