Investigations on Net CO2 Assimilation, Transpiration and Root Growth of Fagus sylvatica Infested with Four Different Phytophthora Species

Abstract: In this contribution, we compare the influence of four different Phytophthora species on root development, net CO₂ assimilation and transpiration of beech seedlings and saplings. Some few days after inoculation, photosynthesis and transpiration of seedlings infected with either P. citricola or P. cambivora were strongly reduced. In parallel, about 60 % of their root systems was destroyed compared to control plants. Three weeks after infection, all seedlings were dead, showing severe wilt symptoms on leaves. Remarkably, P. syringae and P. undulata infected seedlings and older beech plants did not differ from controls regarding photosynthesis and transpiration, although the root systems were damaged. However, a significant influence on net CO₂ assimilation and transpiration of P. citricola infected beech saplings was visible after bud break in the following year. Some days before plants started to wilt, photosynthesis and transpiration were reduced to almost zero. Water use efficiency data (WUE) clearly indicated that infected plants suffered from severe drought.     

Virus Infection of Forest Trees by Mechanical Transmission

Abstract Back transmission trials on young forest plants with isolates of purified viruses from the same tree species were performed using different inoculation techniques. Spruce seedlings and willow plants were successfully infected with tobacco necrosis virus (TNV) by the conventional method of mechanical inoculation of virus suspension mixed with celite as abrasive. Cherry leaf roll virus (CLRV) was transmitted to birches only after adding poly-L-orithine (PLO) to the inoculum. The same method was successful with brome mosaic virus (BMV) on beech seedlings. PLO also improved the rate of infection on TNV in willows. In only one case, was CLRV transmitted conventionally to a white ash seedling. The infection of white ash was increased when frozen powders, of infected ash leaves were directly rubbed onto leaves. BMV could not be transmitted to beech seedlings by carborundum pressure-inoculation. Stem slashing-inoculation of BMV and CLRV was successful with CLRV in one beech out of 60 seedlings.     

Acclimation to ozone affects host/pathogen interaction and competitiveness for nitrogen in juvenile Fagus sylvatica and Picea abies trees infected with Phytophthora citricola

In a two-year phytotron study, juvenile trees of European beech (Fagus sylvatica) and Norway spruce (Picea abies) were grown in mixture under ambient and twice ambient ozone (O3) and infected with the root pathogen Phytophthora citricola. We investigated the influence of O3 on the trees' susceptibility to the root pathogen and assessed, through a 15N-labelling experiment, the impact of both treatments (O3 exposure and infection) on belowground competitiveness. The hypotheses tested were that: (1) both P. citricola and O3 reduce the belowground competitiveness (in view of N acquisition), and (2) that susceptibility to P. citricola infection is reduced through acclimation to enhanced O3 exposure. Belowground competitiveness was quantified via cost/benefit relationships, i.e., the ratio of structural investment in roots relative to their uptake of 15N. Beech had a lower biomass acquisition and captured less 15N under enhanced O3 and P. citricola infection alone than spruce, whereas the latter species appeared to profit from the lower resource acquisition of beech in these treatments. Nevertheless, in the combined treatment, susceptibility to P. citricola of spruce was increased, while beech growth and 15N uptake were not further reduced below the levels found under the single treatments. Potential trade-offs between stress defence, growth performance, and associated nitrogen status are discussed for trees affected through O3 and/or pathogen infection. With respect to growth performance, it is concluded that O3 enhances susceptibility to the pathogen in spruce, but apparently raises the defence capacity in beech..     

Diversity in Pathogenicity to Tobacco and in Elicitin Production among Isolates of Phytophthora parasitica

The pathogenicity to tobacco of a large set of Phytophthora parasitica isolates has been assessed using several procedures: root inoculation of young plants, leaf inoculation on detached disks and stem inoculation of decapitated plants, with or without healing. Analysing various aspects of the plantpathogen interaction with this array of tests led to the discrimination between three groups of isolates. In the isolates from hosts other than tobacco, none was truly pathogenic to tobacco, and all but one produced parasiticein, a proteinaceous elicitor of the elicitin family which induces a hypersensitive-like response in tobacco. Isolates producing parasiticein in vitro induced necrotic fleks on the leaves upon inoculation of roots orof, freshly wounded stems. Most tobacco isolates, including all the highly virulent ones, were characterized by a lack of elicitin production. However, those collected in Australia and Zimbabwe differed in that they exhibited reduced virulence, induced leaf, necrotic flecks and produced parasiticein. The incidence of elicitin production on virulence and the significance of two types of tobacco-pathogenic strains for tobacco pathology are discussed.     

光强对烟草幼苗形态和生理指标的影响

通过白纱布遮荫模拟不同光生境条件(透光率分别为100%、68.2%、35.4%和16.7%),研究了光强因子对烟草幼苗形态和生理指标的影响.结果表明:随相对光强的减弱,幼苗高度增加,茎粗、干鲜比、叶片厚度和单位叶面积质量均呈降低趋势,幼苗干物质积累减少,但其对叶数的影响不大.弱光条件下,叶片自由水、叶绿素、总氮和蛋白质含量增加,束缚水含量降低,叶绿素a/b值减小,转化酶活性降低;烟草幼苗根系相对不发达,根冠比和根生物量减小,根系活力降低.表明弱光条件不利于培育烟草壮苗,生产中应尽可能改善苗床的光照条件.     

Mycorrhizosphere responsiveness to atmospheric ozone and inoculation with Phytophthora citricola in a phytotron experiment with spruce/beech mixed cultures

The aim was to analyze functional changes in the mycorrhizosphere (MR) of juvenile spruce and beech grown in a mixture under ambient and twice ambient ozone and inoculated with the root pathogen Phytophthora citricola. The phytotron experiment was performed over two vegetation periods, adding the pathogen at the end of the first growing season. Root biomass data suggest that the combined treatment affected spruce more than beech and that the negative influence of ozone on stress tolerance against the root pathogen P. citricola was greater for spruce than for beech. In contrast, beech was more affected when the pathogen was the sole stressor. The functional soil parameter chosen for studies of MR soil samples was activity of extracellular enzymes. After the first year of ozone exposure, MR soil samples of both species showed increased activity of almost all measured enzymes (acid phosphatase, chitinase, beta-glucosidase, cellobiohydrolase) in the O3 treatment. Species-specific differences were observed, with a stronger effect of P. citricola on beech MR and a stronger ozone effect on spruce MR. In the second year, the effects of the combined treatment (ozone and P. citricola) were a significant increase in the activity of most enzymes (except cellobiohydrolase) for both tree species. The results indicated that responsiveness of MR soils towards ozone and P. citricola was related to the severity of infection of the plants and the reduction of belowground biomass, suggesting a strong, direct influence of plant stress on MR soil enzyme activity. Additional research is needed using different species and combined stresses to determine the broader ecological relevance of shifts in rhizosphere enzymes.     

Phosphite Protects Fagus sylvatica Seedlings towards Phytophthora plurivora via Local Toxicity,Priming and Facilitation of Pathogen Recognition

Phytophthora plurivora causes severe damage on Fagus sylvatica and is responsible for the extensive decline of European Beech throughout Europe. Unfortunately, no effective treatment against this disease is available. Phosphite (Phi) is known to protect plants against Phytophthora species; however, its mode of action towards P. plurivora is still unknown. To discover the effect of Phi on root infection, leaves were sprayed with Phi and roots were subsequently inoculated with P. plurivora zoospores. Seedling physiology, defense responses, colonization of root tissue by the pathogen and mortality were monitored. Additionally the Phi concentration in roots was quantified. Finally, the effect of Phi on mycelial growth and zoospore formation was recorded. Phi treatment was remarkably efficient in protecting beech against P. plurivora; all Phi treated plants survived infection. Phi treated and infected seedlings showed a strong up-regulation of several defense genes in jasmonate, salicylic acid and ethylene pathways. Moreover, all physiological parameters measured were comparable to control plants. The local Phi concentration detected in roots was high enough to inhibit pathogen growth. Phi treatment alone did not harm seedling physiology or induce defense responses. The up-regulation of defense genes could be explained either by priming or by facilitation of pathogen recognition of the host.     

The biology of Peronospora viciae on pea: factors affecting the susceptibility of plants to local infection and systemic colonization

Peronospora viciae (Berk.) Casp. penetrated leaf disks of Pisum sativum L. through the cuticle. Resistance of pea plants and of individual leaves to infection by P. viciae increased with age, but decreased again at senescence. Resistance was shown by a restriction in fungal growth and sporulation and by a chlorotic reaction in the leaves. Systemic invasion followed infection of meristematic tissue, and was induced by inoculation into the apical bud of young plants, or on to the epicotyl or hypocotyl, but not roots of germinating seedlings. Most plants whose growth was retarded showed an increased resistance to systemic infection. Pods were infected externally by sporangia, rather than by mycelial growth through the peduncle and pedicel. Oospores and mycelium were found in the testas of some seeds, but seeds from infected pods did not give rise to infected seedlings.     

根结线虫对文冠果苗木根系结构与生长及矿质养分分布特性的影响北大核心CSCD

以根结线虫侵染的文冠果一年生苗木为材料,观察分析苗木根系的形态和显微结构,植株生长发育以及主要矿质养分在苗木中的分布特性,并分析不同矿质元素与线虫侵染的关联关系,以揭示根结线虫对文冠果苗木生长发育的影响机制。结果表明:(1)受根结线虫侵染的文冠果苗木根系形成根结,依据根结发生程度分为0(对照,正常植株)、2、3、4级;与对照相比,具有根结的文冠果苗木根系解剖结构特征主要表现为皮层相对较厚,木质部排列扭曲,导管较少,射线数目较少且分布不均匀;韧皮细胞内含物明显较多;多个巨细胞及细胞空腔主要存在于韧皮部。(2)具根结的苗木株高和地径较对照均增加,且株高增幅达显著水平(P<0.05)。(3)具根结苗木根、茎、叶部位的N、P含量均较对照下降,根部降幅达显著水平(P<0.05),其中具2~4级根结的根部N含量分别较对照显著下降7.8%、16.0%和29.5%,P含量分别显著下降15.6%、7.1%和43.3%;根部Fe含量、Zn含量显著上升,2~4级根结根系中Fe含量较对照分别增加1.56倍、0.81倍和3倍,Zn含量分别增加1.11倍、1.56倍和1.78倍;具3和4级根结的苗木根系中K含量较对照显著增加(P<0.05),具2和4级根结的苗木叶片中K含量较对照分别显著下降61.5%和47.0%。(4)苗木根部、茎部N元素含量对于根结线虫侵染的响应最明显,且其含量随侵染程度的增大而降低,叶部Mg元素含量与根结线虫侵染率、K元素含量与整株苗木生物量均呈显著负相关关系(P<0.05)。该研究为根结线虫对文冠果苗木养分吸收利用及运转的影响提供了一定的理论依据。     

The influence of present-day levels of ultraviolet-B radiation on seedlings of two Southern Hemisphere temperate tree species

Seedlings of two Southern Hemisphere temperate trees species (mountain beech: Nothofagus solandri var. cliffortioides (Hook. f.) Poole and broadleaf: Griselinia littoralis Raoul) were grown in the field to determine the effects of present-day levels of ultraviolet-B radiation (UV-B) on growth, biomass, UV-B absorbing compounds, leaf optical properties and photoinhibition. Plants were covered with either UV-B transmitting or UV-B absorbing filters. After 125 days of typical summer weather, total biomass of both species was not affected by the UV-B treatments. Without UV-B, height increased (23%) and the number of leaves produced decreased (–21%) in beech, but broadleaf was unaffected. The effect of UV-B on beech height and leaf number was manifest during a second flush of leaves suggesting differences in response to UV-B of leaves initiated in different seasons and UV-B radiation regimes. Leaves of both species were essentially opaque to the transmission of UV-B. In the absence of UV-B the transmission of photosynthetically active radiation through leaves of both species increased, foliar nitrogen concentrations increased and levels of UV-B absorbing compounds decreased. In the youngest leaves of beech but not of broadleaf, removal of UV-B reduced midday photoinhibition, and did not alter the complete recovery of the fluorescence ratio F_V/F_M in the evening to predawn levels. As leaves of both species aged, midday photoinhibition decreased, with the result that UV-B had no effect on photoinhibition in mature leaves. Results of this experiment show that even under present-day UV-B levels, UV-B radiation modifies the physiology, optical properties and secondary compounds of leaves of both beech and broadleaf seedlings.     

Does water and phosphorus uptake limit leaf growth of Rhizoctonia-infected wheat seedlings?

Wheat seedlings infected with a pure inoculum of the root-rotting fungus Rhizoctonia solani were grown in pots designed to fit in pressure chambers, to allow the effects of the Rhizoctonia infection on leaf growth to be studied while maintaining the leaves at elevated water status. Wheat was grown to the third leaf stage in soil inoculated with three different levels of Rhizoctonia, and the pots were then pressurised for seven days to maintain the leaf xylem at the point of bleeding (ie. the leaves were at full turgor). The reduction in leaf expansion caused by Rhizoctonia was not overcome by pressurisation, indicating that a reduced supply of water to the leaves was not responsible for reduced leaf growth. The addition of phosphorus to pots marginally deficient in P did not increase the leaf growth of Rhizoctonia-infected plants, despite increased P uptake to the leaves. These results indicate that a reduced supply of water to the leaves and a supply of phosphorus that was bordering on deficient was not the cause of the growth reduction in seedlings with Rhizoctonia infection. The nature of this reduced growth remains uncertain but may involve growth regulators produced by the fungus, or by the plant as a result of the infection process. The mechanism of these growth reductions is of interest as it may provide a key to the development of plant resistance mechanisms. This revised version was published online in June 2006 with corrections to the Cover Date.     

SOME EFFECTS OF VIRUS INFECTION ON LEAF WATER CONTENTS OF NICOTIANA SPECIES

Infection with tobacco mosaic virus decreases the water content which detached tobacco leaves attain when kept for 20 hr. in conditions of minimum water stress, and does so more when the plants are kept in light before inoculation than when they are kept in darkness. No such effects of infection during the first day after inoculation were obtained with tobacco leaves infected with either tobacco etch virus or potato virus X , or with Nicotiana glutinosa leaves infected with tobacco mosaic virus. These results, like those showing early effects of TMV on respiration and photosynthesis of tobacco leaves, suggest that inoculation with TMV affects deeper leaf tissues than the epidermis earlier in tobacco leaves than in other leaves, and earlier than other viruses in tobacco leaves.     

Trypsin inhibitor activity in mature tobacco and tomato plants is mainly induced locally in response to insect attack,wounding and virus infection

Wounding of plants by insects is often mimicked in the laboratory by mechanical means such as cutting or crushing, and has not been compared directly with other forms of biotic stress such as virus infection. To compare the response of plants to these types of biotic and abiotic stress, trypsin inhibitor (TI) activity induced locally and systemically in mature tobacco (Nicotiana tabacum L.) and tomato (Lycopersicon esculentum L.) plants was followed for 12 days. In tobacco, cutting, crushing and insect feeding all induced comparable levels of TI activity of approx. 5 nmol·(mg leaf protein)^?1 in wounded leaves, while tobacco mosaic virus (TMV) infection of tobacco induced 10-fold lower amounts in the infected leaves. In tomato, feeding by insects also led to the induction of a level of TI activity of 5 nmol·(mg leaf protein)^?1. In contrast, both cutting and crushing of tomato leaves induced 10-fold higher amounts. These data show that biotic stress, in the form of insect feeding and TMV infection, and abiotic stress, in the form of wounding, have different effects on local levels of induced TI activity in mature tobacco and tomato plants. Irrespective of the type of wounding, in neither tobacco nor tomato could systemic induction of TI activity be observed in nearby unwounded leaves, which suggests that systemic induction of TI activity in mature tobacco and tomato plants is different from systemic TI induction in seedlings. Wounding of tobacco leaves, however, did increase the responsiveness to wounding elsewhere in the plant, as measured by an increased induction of TI activity.     

Fulvic Acid,Brassinolide, and Uniconazole Mediated Regulation of Morphological and Physiological Traits in Maize Seedlings Under Water Stress

Water stress is one of the most important factors limiting sustainable crop production. Therefore, the effects of the plant growth regulators (PGRs) fulvic acid (FA), brassinolide (BR), and uniconazole (Uni) on seedling growth and physiology of two maize (Zea mays L.) varieties were evaluated under???0.7 MPa water stress induced by polyethylene glycol-6000. Under drought stress, the PGRs promoted seedling growth, altered the root-to-shoot ratio, and significantly increased root biomass, length, surface area, diameter, and volume. In addition, depending on the PGR, net photosynthesis rate, SPAD value (indicating chlorophyll content), and water use efficiency increased significantly, under drought stress, whereas transpiration rate decreased. The PGRs also significantly increased antioxidant enzyme activities and significantly decreased malondialdehyde accumulation in leaves and roots under drought stress. Zhengdan958 showed greater variation in physiological responses and stronger drought resistance than Xundan20. In alleviating drought stress in maize seedlings, FA had the greatest effects on shoot growth and leaf physiology; Uni exerted its effects by regulating root structure, and BR effects were intermediate. Under drought stress, the three PGRs increased maize seedling growth, which reduced drought stress-induced damage and improved plant ability to resist the adversity. Based on a comprehensive analysis of physiological indices of drought resistance, Uni is recommended as the best PGR to improve maize seedlings resistance to drought.

     

Photosynthetic Responses of a Temperate Liana to Xylella fastidiosa Infection and Water Stress

Xylella fastidiosa is a xylem‐limited bacterial plant pathogen that causes bacterial leaf scorch in its hosts. Our previous work showed that water stress enhances leaf scorch symptom severity and progression along the stem of a liana, Parthenocissus quinquefolia, infected by X. fastidiosa. This paper explores the photosynthetic gas exchange responses of P. quinquefolia, with the aim to elucidate mechanisms behind disease expression and its interaction with water stress. We used a 2 × 2‐complete factorial design, repeated over two growing seasons, with high and low soil moisture levels and infected and non‐infected plants. In both years, low soil moisture levels reduced leaf water potentials, net photosynthesis and stomatal conductance at all leaf positions, while X. fastidiosa‐infection reduced these parameters at basally located leaves only. Intercellular CO₂ concentrations were reduced in apical leaves, but increased at the most basal leaf location, implicating a non‐stomatal reduction of photosynthesis in leaves showing the greatest disease development. This result was supported by measured reductions in photosynthetic rates of basal leaves at high CO₂ concentrations, where stomatal limitation was eliminated. Repeated measurements over the summer of 2000 showed that the effects of water stress and infection were progressive over time, reaching their greatest extent in September. By reducing stomatal conductances at moderate levels of water stress, P. quinquefolia maintained relatively high leaf water potentials and delayed the onset of photosynthetic damage due to pathogen and drought‐induced water stress. In addition, chlorophyll fluorescence measurements showed that P. quinquefolia has an efficient means of dissipating excess light energy that protects the photosynthetic machinery of leaves from irreversible photoinhibitory damage that may occur during stress‐induced stomatal limitation of photosynthesis. However, severe stress induced by disease and drought eventually led to non‐stomatal decreases in photosynthesis associated with leaf senescence.     

等渗的盐分和水分胁迫对芦荟幼苗生长和离子分布的效应

 研究了等渗透势（-0.44、-0.88 MPa）NaCl和PEG 6000处理对六叶龄芦荟(Aloe vera)幼苗叶片生长速率、干物质积累、电解质渗漏和离子吸收、分配的效应。结果表明: -0.44、-0.88 MPa NaCl和PEG处理10 d均明显抑制芦荟幼苗叶片伸长生长,植株干物质积累速率显著降低, 叶片含水量降低,叶片细胞电解质渗漏率上升。NaCl对芦荟幼苗生长的抑制作用显著大于PEG处理的。不同器官离子含量、根系和叶片横切面X-射线微区分析结果表明, NaCl胁迫导致芦荟体内Na+、Cl－含量显著上升,根中增幅明显高于叶片,其中Cl－尤为显著。NaCl胁迫严重抑制芦荟对K+ 和Ca2+ 的吸收及其向叶片的运输,根、叶K+/Na+、Ca2+/Na+ 比率显著下降,而PEG胁迫对离子平衡的干扰较轻,是芦荟对水分胁迫的适应能力高于盐胁迫的主要原因之一。但芦荟对 -0.44～-0.88 MPa NaCl胁迫仍有一定的适应能力,主要原因是：1) 根系对离子的选择性吸收和运输较强,并随着盐胁迫强度增加其选择性增强; 2) 芦荟叶片中的盐分在贮水组织中显著积累,明显高于其它组织细胞。同时,芦荟是CAM（景天酸代谢）途径植物,蒸腾极小,盐分随蒸腾流进入地上部的机会小。     

Elicitin 172 from an isolate of Phytophthora nicotianae pathogenic to tomato

Elicitin 172, an acid protein with elicitor activity, has been isolated in true form from culture filtrates of Phytophthora nicotianae, the causal agent of crown and root rot of tomato (Lycopersicon esculentum). The M(r) (10,349 +/- 1) of the purified protein, determined by ES-MS, is identical to that calculated for parasiticein using the mean isotopic composition and assuming the occurrence of three disulfide bridges. The primary structure of elicitin 172, determined using also MALDI-MS experiments, shows complete identity with parasiticein, with elicitin 310 and a cloned elicitin gene from P. parasitica (= P. nicotianae), confirming conservation of the elicitin sequence within a single species. The protein induces necrosis (hypersensitive reaction) on tobacco, but no symptoms on tomato, when applied on the leaves. Tomato pretreated with elicitin 172 was affected by P. nicotianae, as well as by the phytotoxic aggregates, naturally occurring with the elicitin in the non permeated dialysis fraction of culture filtrates. Finally, the elicitin induce protection of capsicum (Capsicum annuum) and vegetable marrow (Cucurbita pepo) from P. capsici.     

Effects of ozone and Phytophthora citricola on non-structural carbohydrates of European beech (Fagus sylvatica) saplings

A lysimeter study was performed to monitor long term effects of chronic ozone enrichment on saplings of European beech (Fagus sylvatica L). After 3 years of ozone exposure a root infection with Phytophthora citricola Swada was established in the fourth year to study the interaction between elevated ozone and the root infection on the carbon budget of beech saplings. By using quantitative PCR no differences in root infection with P. citricola were observed between the ozone treatments. In contrast to the first 3 years of ozone exposure, sucrose and starch concentrations in leaves were diminished in ozone treated plants in the fourth year. The root infection reduced sucrose concentrations in leaves. Starch reserves of the heterotrophic biomass were not affected by any treatments. Thus 4 years of ozone exposure and 1 year of P. citricola root infection had only limited effect on carbohydrate metabolism in beech saplings.     

The Role of Infective Plant Debris, and its Concentration in Soil, in the Ecology of Tomato Mosaic Tobamovirus—a Non-vectored Plant Virus

Only a few plants in a crop are generally thought to become infected by abiotic soil transmission. In glasshouse experiments we have induced almost total infection in tomatoes growing in soil with infective debris, but levels of infection are dependent upon certain conditions. We found that as the inoculum concentration decreased (i) a greater percentage of the infections were either latent (symptomless) or restricted to roots and (ii) infection levels decreased. Thus, apparent infection (based on symptoms) of 0. 10% was in reality 0 60–70% based on enzyme-linked immunosorbent assay and immunoelectron microscopy testing of roots and leaves. This occurred whether seedlings or seed were planted into infected mix. Under conditions in which minimal root damage was caused (planting seed) or roots were mechanically inoculated only once, almost all systemically infected plants were symptomless.     

AN ANALYSIS OF THE GROWTH RESPONSE OF YOUNG TOMATO PLANTS TO INFECTION BY VERTICILLIUM ALBO-ATRUM

Infection of seedling tomatoes with Verticillium albo-atrum checked growth but did not result immediately in leaf yellowing. Localized wilting occurred in some plants 2 weeks after the check to growth was evident. 8 weeks after inoculation, dry weights of leaf, stem and root were decreased by 72, 70, and 65% respectively.
Of the growth attributes studied, leaf area was most reduced by infection and this was due to a failure of the leaves to expand rather than to a fall in the rate of leaf production. Neither water nor nitrogen appeared to be limiting factors in this respect. The water content of infected leaves was not reduced until 6 weeks after inoculation, when leaf yellowing and necrosis had also appeared. The percentage N contents of stem, root and leaf of infected plants exceeded those of the healthy controls 24 days after inoculation. N uptake was not seriously impaired until 21 days later.
The photosynthetic efficiency of the green leaves of infected plants was reduced. The mean values for net assimilation rates were: Healthy 0.47 and infected 0.39 g./dm.²/week.
Plants, in which two-thirds of the root system had been killed by crushing, were placed in contact with mycelium in soil. This initial root injury did not significantly affect the growth of infected plants.
The data accord with a toxin theory of damage to infected plants, but the slow development of chlorosis and wilting symptoms in the young plants suggested a greater tolerance to the toxin than is found in older plants.