Evidence for Phytophthora cinnamomi involvement in Iberian oak decline

Rapid and sometimes extensive mortality and decline of oak, principally Quercus suber and Q. ilex , has occurred in parts of southern Spain and Portugal in recent decades. We report here isolation of the aggressive root pathogen Phytophthora cinnamomi from roots of diseased oaks or from soil at eleven out of thirteen decline foci examined. It is proposed that the introduction and spread of P. cinnamomi may be a major factor in the Iberian oak decline, interacting with drought and other site factors, and leading to stress-related attacks by insects and other fungi. By analogy, it may also be involved in the similar oak declines occurring elsewhere on the Mediterranean.     

Seasonal changes in susceptibility of Quercus suber to Botryosphaeria stevensii and Phytophthora cinnamomi

Monthly inoculations of both intact plants and excised shoots of Quercus suber with the pathogenic species Botryosphaeria stevensii and Phytophthora cinnamomi were performed to investigate seasonal changes in susceptibility of this forest tree species in relation to environmental parameters and plant water status. Infection symptoms were mainly detected on seedlings inoculated from spring to autumn (April through October) with either pathogen. Mean canker sizes also showed a seasonal pattern, the higher values being recorded in the same period as above. Lesion lengths were significantly ( P  < 0·001) related to environmental minimum temperature. Mean daily minimum temperatures within the range of 5–12°C clearly inhibited lesion development of P. cinnamomi , whereas B. stevensii showed a less pronounced decrease in canker expansion at the same temperature range. In excised shoots of Q. suber inoculated monthly with B. stevensii , a negative linear relationship was found between the studied range of plant relative water content (81–91%) and canker length. In contrast, the lesions caused by P. cinnamomi were not significantly ( P  = 0·32) related to any seasonal change in water content. Some control measures for the diseases caused by both pathogens are discussed on the basis of the seasonal changes in host susceptibility observed in this study.     

Testing variability in pathogenicity of Phytophthora cinnamomi

Forty eight isolates of Phytophthora cinnamomi from various host plants in France (35 isolates) and in other countries were tested for pathogenicity. Seedlings of chestnut, northern red oak, pine and eucalyptus were infected by soil contamination. Taproots, stems and bark strips of plants of chestnut and different oak species were inoculated with mycelium agar disks. Results of the different experiments were in good agreement. All isolates appeared pathogenic to all the different test species but with variable levels of virulence. Isolates with consistent low or high level of virulence, which could be used as standards in further studies, were identified. Interaction between P. cinnamomi isolates and host plant species was significant in terms of lesion lengths. These interactions could not be related to host from which P. cinnamomi was isolated. Consistent with this, in Quercus rubra, the isolate-provenance interaction was not significant. This feature is encouraging for provenance screening for resistance to P. cinnamomi in this species. The variation in virulence was not related to other isolate characteristics (mating type, electrophoretic type, age).     

In vitro leaf inoculation studies as an indication of tree foliage susceptibility to Phytophthora ramorum in the UK

Leaves of 11 coniferous and 23 broad-leaved tree species important to UK forestry were tested for their susceptibility to the quarantine pathogen Phytophthora ramorum using a detached leaf assay. Two European and two USA isolates were used. Wounded and unwounded leaves were dipped in zoospore suspensions during summer; conifers were also tested in winter. Successful infection of tissue and amount of necrosis were assessed. Highly susceptible broad-leaved hosts included Aesculus hippocastanum , Fraxinus excelsior , Quercus ilex , Ulmus procera and, to a lesser extent, Castanea sativa , Q. cerris and Q. petraea , together with Umbellularia californica and rhododendrons. Acer pseudoplatanus , Alnus glutinosa , Carpinus betulus , Corylus avellana , Fagus sylvatica , Prunus avium , Q. robur , Q. rubra and Q. suber showed consistently low susceptibility. Conifer species including Abies procera , Picea abies , P. sitchensis , Pseudotsuga menziesii , Sequoia sempervirens and Tsuga heterophylla were also susceptible. Pseudotsuga menziesii and A. procera were severely affected. Pinus contorta , P. nigra var. maritima and P. sylvestris were virtually resistant, while Taxus baccata was only slightly affected. Increased necrosis was apparent on leaves that were wounded prior to inoculation. These results extend the known range of trees that P. ramorum is able to attack and confirm its relative host-nonspecificity.     

Susceptibility of Iberian trees to Phytophthora ramorum and P. cinnamomi

The capacity of Phytophthora ramorum to colonize the inner bark of 18 native and two exotic tree species from the Iberian Peninsula was tested. Living logs were wound-inoculated in a growth chamber with three isolates belonging to the EU1 and two to the NA1 clonal lineages of P. ramorum . Most of the Quercus species ranked as highly susceptible in experiments carried out in summer, with mean lesion areas over 100 cm² in Q. pubescens , Q. pyrenaica , Q. faginea and Q. suber and as large as 273 cm² in Q. canariensis , ca . 40 days after inoculation. Quercus ilex ranked as moderately susceptible to P. ramorum , forming lesions up to 133 cm² (average 17·2 cm²). Pinus halepensis and P. pinea were highly susceptible, exhibiting long, narrow lesions; but three other pine species, P. pinaster , P. nigra and P. sylvestris , were resistant to slightly susceptible. No significant difference in aggressiveness was found between the isolates of P. ramorum . In addition, there was evidence of genetic variation in susceptibility within host populations, and of significant seasonal variation in host susceptibility in some Quercus species. The results suggest a high risk of some Iberian oaks to P. ramorum , especially in forest ecosystems in southwestern Spain, where relict populations of Q. canariensis grow amongst susceptible understory species such as Rhododendron ponticum and Viburnum tinus . One isolate of P. cinnamomi used as positive control in all the inoculations was also highly aggressive to Iberian oaks and Eucalyptus dalrympleana .     

European oak declines and global warming: a theoretical assessment with special reference to the activity of Phytophthora cinnamomi

Causes of current severe declines of the deciduous oaks Quercus robur and Q. petraea in northern and central Europe and of the evergreen Q. ilex, Q. suber and other Quercus spp. in the Mediterranean area are reviewed. Factors implicated include drought, pollution, winter cold, flooding, and stress-related attacks by insects and fungi. Additional factors in Mediterranean oak declines include changing land-use patterns and root disease caused by the aggressive, exotic oomycete root pathogen Phytophthora cinnamomi. Under conditions of global warming the survival and degree of root disease caused by this fungus seems likely to be enhanced, while the host range of the organism might also be increased. Application of the CLIMEX climate-matching program suggests that with a mean increase in temperatures of 1.5–3°C the fungus might considerably increase its disease activity in its existing locations, and to some extent spread northwards and eastwards. However, it seems unlikely to become significantly active in areas of Europe with colder winters such as parts of Scandinavia, Russia and the central Danube. The predictive value of research on major environmental problems such as oak declines could be enhanced by more highly coordinated European forestry research programmes.     

Phytophthora species in oak ecosystems in Turkey and their association with declining oak trees

From 1999 to 2001, a survey on the occurrence of Phytophthora spp. in the rhizosphere soil of healthy and declining oak trees was conducted in 51 oak stands in Turkey. Seven Phytophthora spp. were recovered from six out of the nine oak species sampled: P .  cinnamomi , P .  citricola , P .  cryptogea , P .  gonapodyides , P .  quercina , Phytophthora sp. 1 and Phytophthora sp. 2. The most frequently isolated species, P .  quercina , was very common on slopes susceptible to drought. It occurred in four different climatic zones and on six Quercus spp., suggesting that it is native to oaks. The second most common species, P .  citricola , was separated into three subgroups: type C was recovered only in Anatolia, whereas A and B occurred only in the European part of Turkey. Phytophthora cinnamomi was recovered at one site only, and may not be involved in oak decline in Turkey. The other four species were recovered sporadically. On affected sites there was a significant association between deteriorating crown status and the presence of Phytophthora spp., particularly P .  quercina . The occurrence of Phytophthora species was significantly influenced by soil pH. Stem inoculation tests on oak seedlings revealed that Q .  petraea was the most susceptible species.     

Cryptogein and capsicein promote defence responses in Quercus suber against Phytophthora cinnamomi infection

The decline of cork oak (Quercus suber) stands in Iberian Peninsula is associated with infection by Phytophthora cinnamomi. Most Phytophthora species secrete elicitins, which can enhance defence reactions against some pathogens. Here cytological and physiological effects of the elicitins cryptogein and capsicein on cork oak root infection by P. cinnamomi were evaluated. The progression of the pathogen in root tissue and its effects on total fatty acid (TFA) composition of roots and leaves were analysed in seedlings. Net photosynthesis (P _n ), stomatal conductance (g _s ), chlorophyll a fluorescence measurements (quantum yield of linear electron transport ? _e , photochemical quenching q _P, non-photochemical quenching NPQ) and carotenoid determinations were carried out in well established (4?months) plants. In elicitin-treated roots, 2?days after inoculation, the pathogen which presented loss of viability and membrane degradation was mainly restricted to the intercellular spaces of the cortical parenchyma, and did not reach the vascular cylinder. Electron dense materials accumulated in the intercellular spaces of the cortex next to disorganized hyphae, suggested to be related with defence reactions. Cryptogein (or its interaction with P. cinnamomi) induced enhanced lipid synthesis in leaves, which may contribute to preserve membrane stability. P. cinnamomi decreased P _n , g _s , ? _e , and q _P, whereas elicitin-treated plants displayed values similar to controls. Overall, the results indicated a resistance response of cork oak against this oomycete, induced by the elicitins.     

Comparison of the Susceptibility of Quercus petraea, Q. robur and Q. rubra to Collybia fusipes

Collybia fusipes is the cause of a root rot of Quercus petraea (sessile oak), Q. robur (pedunculate oak) and Q. rubra (red oak). This parasite is often reported to cause problems in stands of red oaks and field evidence suggests that this North American oak species is more susceptible than the two European oak species. Young saplings of the three oak species and of chestnut, Castanea sativa, in the glasshouse, and also mature sessile and red oaks growing in the same stand were inoculated with C. fusipes to compare their susceptibility. Red oak, both as young seedlings and mature trees, was more susceptible to C. fusipes than sessile oak. Chestnut seedlings were as susceptible as sessile oak. Susceptibility of pedunculate oak seedlings was intermediate between red oak and sessile oak. In one experiment this species was significantly less susceptible than red oak, and in the other it was as susceptible.     

Effect of <Emphasis Type="Italic">Meloidogyne ethiopica</Emphasis> parasitism on water management and physiological stress in tomato

Root-knot nematodes (RKN) are obligate endoparasites that severely damage the host root system. Nutrient and water uptake are substantially reduced in infested plants, resulting into altered physiological processes and reduced plant growth. The effect of nematode infestation on the morphological changes of roots and subsequent physiological plant responses of infested tomatoes with the RKN Meloidogyne ethiopica was studied in a pot experiment. Plants were infested with two inoculum densities (10 or 50 eggs per cm³ substrate) and its effect was evaluated 74 and 102 days post inoculation (DPI). Morphological changes and root growth was determined by analysing scanned images of the whole root system. Nematode infestation reduced the portion of fine roots and increased that of coarse roots due to gall formation. Fine roots of non-infested control plants represented around 51% of the area of the whole root system at 74 and 102 DPI. In comparison to controls, plants inoculated with low and high nematode density had 2.1 and 3.2-times lower surface area of fine roots at 102 DPI. Root analyses revealed that plants had a very limited ability to mitigate the effects of the root-knot nematodes infestation by altering root growth. Root galls had a major influence on the hydraulic conductivity of the root system, which was significantly reduced. The low leaf water potential of infested plants coincided with decreased stomatal conductivity, transpiration and photosynthesis. The latter two were reduced by 60–70% when compared to non-infested control plants.     

Anti-Phytophthora cinnamomi activity of Phlomis purpurea plant and root extracts

Phlomis purpurea (Lamiaceae), found in Quercus suber and Quercus ilex ssp. rotundifolia forest habitats in southern Portugal, is a non-host for the oomycete Phytophthora cinnamomi, the main biotic factor involved in cork oak and holm oak decline in the Iberian Peninsula. The effect of P. purpurea crude ethanol root extract was evaluated in vitro on P. cinnamomi mycelial growth, sporangial production, zoospore release and germination as well as on chlamydospore production and viability. The protection of cork oak against infection by the pathogen was also evaluated in planta. At 10 mg ml^?1, in vitro inhibition of the pathogen structures was 85–100 %. In addition, P. purpurea plants were shown to protect Q. suber and Q. ilex from P. cinnamomi infection and to reduce the inoculum potential in glasshouse trials, indicating the ability to reduce root infection by the pathogen. The results suggest that P. purpurea has the potential to reduce disease spread and that their root extracts could provide candidate substances for control of the important pathogen, P. cinnamomi.     

不同土壤水分含量对玉米气孔发育过程和蒸腾耗水量的影响

以盆栽玉米为材料,采用三个水分梯度进行处理,显微观察和比较了新生叶的气孔密度、气孔器的长度和宽度、气孔开张度,测定了水分胁迫对气孔导度、植株蒸腾耗水量和各个器官生物量的影响.结果表明:土壤水分含量降低使得玉米叶片气孔密度增大,气孔导度减小,气孔器宽度的降低幅度明显大于长度的降低.轻度和中度土壤水分胁迫使得全生育期单株蒸...     

Hydraulic resistance partitioning between shoot and root system and plant water status of Haloxyolon ammodendron growing at sites of contrasting soil texture

Hydraulic resistance components and water relations were studied on Haloxyolon ammoden-dron,a small xeric tree,growing at sites significantly differed in soil texture.Soil water content,leaf water potential(ψl),xylem water potential(ψx),root water potential(ψroot),leaf transpiration rate(TR) and stomatal conductance(gs) were measured at the two sites during the growing season of 2005 and 2006.Leaf spe-cific hydraulic resistance(Rplant) during the whole growing season,hydraulic resistance of plants(Rp),shoots(Rshoot) and roots(Rroot) in the August of both years were calculated and expressed on leaf area basis.The results showed the proportion of the hydraulic resistance of the aerial part(Rshoot) to the Rp was the same to the proportion of the hydraulic resistance of the soil part(Rroot) to the Rp,indicating that both parts were equivalent important to plant water hydraulic system from soil to leaf.Positive significant corre-lations were found between Rp and Rroot,suggesting that root hydraulics resistance was a major determinant of plant hydraulic resistance(Rp) and transpiration rate.The integrated effect of stomatal control,hy-draulic regulation and morphology adjustment enabled plants at heavy soil site surviving the extreme water deficit period.     

Prediction and Mapping of the Impact of Winter Temperature on the Development of Phytophthora cinnamomi-Induced Cankers on Red and Pedunculate Oak in France

ABSTRACT Phytophthora cinnamomi is the causal agent of a perennial canker that develops on the lower bole on northern red oak and pedunculate oak. The disease has a limited range in Europe, being reported only in southwest France. This limited distribution is probably linked to the susceptibility of P. cinnamomi to frost. A model was developed in previous work to estimate the impact of temperatures of <0 degrees C on the winter survival of P. cinnamomi in trunk cortical tissues and on the subsequent development of cankers. In this article, we report the use of this model to simulate canker development in 503 locations across France during a 30-year period. The predicted canker extension decreased sharply when the median P. cinnamomi winter survival index decreased from 0.95 to 0.65, with cankers that poorly developed when the median survival index was lower than 0.5 to 0.6. The actual incidence of the disease in 192 stands located across southwest France was compared with that of the model outputs. Both presence of disease in stands and frequency of cankered trees in infected stands, but not canker size on infected trees, were strongly related to the median P. cinnamomi survival index. No disease was present in stands with median survival index lower than 0.65, and the frequency of cankered trees in infected stands remained very low in stands with a median survival index between 0.65 and 0.70. Aspect was an additional factor explaining disease incidence, while the effect of elevation was likely due to its effect on winter temperatures. Maps of winter suitability to P. cinnamomi-induced cankers on oaks in France are presented.     

Isolation and Characterization of Brenneria quercina, Causal Agent for Bark Canker and Drippy Nut of Quercus spp. in Spain

ABSTRACT The drippy nut disease of oak was first described in California in 1967 and, since then, the causal agent has not been reported in any other area. This study describes for the first time in Europe the isolation of Brenneria (Erwinia) quercina from bark canker in addition to drippy bud and drippy nut in Quercus ilex and Q. pyrenaica. The bark canker and drippy bud symptoms were not previously described as caused by this bacterium. No fungal pathogens were associated with any of the symptoms. Physiological and biochemical characterization identified the pathogenic isolates from Spain as belonging to B. quercina, similar to the reference strain CFBP 1266. Fatty acid profiles of the Spanish isolates also were similar to the strain of B. quercina from California. Serological analysis by indirect immunofluorescence and enzyme-linked immunosorbent assay using polyclonal antisera against the reference strain of B. quercina and one Spanish oak isolate revealed some antigenic heterogeneity between isolates of different origins. Pathogenicity tests demonstrated that the Spanish isolates were able to reproduce internal symptoms of necrosis and acorn exudation in Q. ilex and Q. pyrenaica and suggest that B. quercina may be associated, among other causes, with the oak decline syndrome affecting Spanish oak forests.     

The early development of disease caused by Phytophthora cinnamomi in Eucalyptus marginata and Eucalyptus calophylla grooving in rehabilitated bauxite mined areas

The soil-borne plant pathogen Phytophthora cinnamomi is widely distributed in the jarrah ( Eucalyptus marginata ) forest of Western Australia. Infested areas of the forest are mined for bauxite and the presence of the pathogen could after the survival of trees re-established after mining. Monitoring of 21 revegetated bauxite mined areas found that survival of jarrah and marri ( Eucalyptus calophylla ) trees was high (85–92% and 93–99%, respectively) after 5–7 years but P. cinnamomi was recovered from dead trees. To identify trees for more detailed study, plant symptoms of stress such as suppressed growth, wilting, yellowing of crown, coppice and epicormic growth and visible stem lesions were used. Over a period of 15 months, 30 E. marginata and 28 E. calophylla were carefully excavated and examined for lesions and the presence of P. cinnamomi. P. cinnamomi was consistently isolated from the lignotuber and collar regions of both hosts but never from the roots alone, except in one instance from E. calophylla where it was isolated from a non-lesioned root. In E. calophylla , the lignotuber appears to be very susceptible to invasion by P. cinnamomi in contrast to the roots which appear resistant. The invasion of the pathogen into the lignotuber and collar regions of both species was consistently associated with ponding of water around the plants. This ponding persists for many hours to days after rain and appears to provide an infection court for P. cinnamomi. Development of rehabilitation procedures to reduce this ponding will minimize the risk of tree deaths caused by this pathogen.     

Phytophthora ramorum and Phytophthora kernoviae on naturally infected asymptomatic foliage

Phytophthora ramorum and Phytophthora kernoviae are recently discovered invasive Phytophthoras causing leaf necrosis and shoot tip dieback mostly on ornamental and forest understorey species, but also cause bleeding cankers on stems of a wide range of tree species. Sporulation occurs only on infected shoots or fruits and foliage so foliar hosts are central to the disease epidemiology. In field trials to assess infection in trap plants exposed to natural inoculum of P. ramorum and P. kernoviae on rhododendron in south west England, it was discovered that leaves of the trap plants ( Rhododendron 'Cunninghams White') and holm oak ( Quercus ilex ) were asymptomatically infected and supported sporulation of both pathogens. More than half the rhododendron trap plants exposed to inoculum of P. kernoviae became infected compared with approximately a third of those exposed to P. ramorum in a natural situation. Approximately one third of the infections were detected from asymptomatic foliage for both pathogens. The significance of these findings for plant health regulation based on visual inspection as a measure to prevent introduction and dissemination of both these pathogens is explored and research gaps identified.     

水分与氮素营养对辽春10号小麦旗叶光合作用的影响

在盆栽条件下,研究不同水分条件下氮素营养对辽春10号春小麦籽粒灌浆期旗叶光合特性的影响.结果表明,随着土壤水分状况的改善,春小麦的光合作用不断增强.土壤相对合水量低于45%时,施氮处理春小麦旗叶气孔导度、蒸腾速率和WUEleaf降低,胞间二氧化碳浓度升高,光合速率受非气孔因素制约而降低.当土壤相对含水量在45%～75%时,随着氮肥施用量的增加和土壤相对含水量的提高,春小麦旗叶的光合效率、气孔导度、蒸腾速率和WUEleaf不断增加,胞间二氧化碳浓度逐渐减小,气孔因素制约了光合效率的提高.当土壤相对含水量高于75%时,氮素营养对春小麦旗叶光合的促进作用减弱.     

Gas exchange of Populus euphratica leaves in a riparian zone

Riparian vegetation belts in arid regions of Central Asia are endangered to lose their ecosystem services due to intensified land use.For the development of sustained land use,management knowledge of plant performance in relation to resource supply is needed.We estimated productivity related functional traits at the edges of the habitat of Populus euphratica Oliv.Specific leaf area(SLA) and carbon/nitrogen(C/N) ratio of P.euphratica leaves growing near a former river bank and close to moving sand dunes in the Ebinur Lake National Nature Reserve in Xinjiang,Northwest China(near Kazakhstan) were determined and daily courses of CO2 net assimilation(PN),transpiration(E),and stomatal conductance(gs) of two consecutive seasons were measured during July-August 2007 and June-July 2008.Groundwater level was high(1.5-2.5 m below ground) throughout the years and no flooding occurred at the two tree stands.SLA was slightly lower near the desert than at the former river bank and leaves contained less N in relation to C.Highest E and g s of P.euphratica were reached in the morning before noon on both stands and a second low maximum occurred in the afternoon despite of the unchanged high levels of air to leaf water vapor pressure deficit(ALVPD).Decline of g s in P.euphratica was followed by decrease of E.Water use efficiency(WUE) of leaves near the desert were higher in the morning and the evening,in contrast to leaves from the former river bank that maintained an almost stable level throughout the day.High light compensation points and high light saturation levels of PN indicated the characteristics of leaves well-adapted to intensive irradiation at both stands.In general,leaves of P.euphratica decreased their g s beyond 20 Pa/kPa ALVPD in order to limit water losses.Decrease of E did not occur in both stands until 40 Pa/kPa ALVPD was reached.Full stomatal closure of P.euphratica was achieved at 60 Pa/kPa ALVPD in both stands.E through the leaf surface amounted up to 30% of the highest E rates,indicating dependence on water recharge from the ground despite of obviously closed stomata.A distinct leaf surface temperature(Tleaf) threshold of around 30℃ also existed before stomata started to close.Generally,the differences in gas exchange between both stands were small,which led to the conclusion that micro-climatic constraints to E and photosynthesis were not the major factors for declining tree density with increasing distance from the river.     

不同土壤水分条件下施肥方式对甘薯叶片气体交换的调节作用

采用盆栽试验,以甘薯品种秦薯4号和619为材料,研究了不同土壤水分条件下施肥方式对甘薯叶片气体交换的影响.结果表明,土壤水分和施肥方式对甘薯叶片净光合速率(Pn)、蒸腾速率(Tr)、水分利用效率(WUE)、气孔导度(Gs)、胞间CO2浓度(Ci)以及气孔限制值(Ls)有显著或极显著的影响,水肥之间存在明显的互作效应.与土壤湿润( 田间持水量70%～75%)相比,土壤轻旱(田间持水量40%～45%)对甘薯叶片Pn、Tr、WUE 、Gs、Ci以及Ls的影响较小,而重旱(田间持水量30%～35%)下,Pn、Tr、WUE、Gs和Gs 大幅下降,Ci明显上升.施肥有助于促进提高Pn、Tr、Gs以及Ls,降低Ci,而且氮磷配施的效果优于氮肥单施.Ci和Ls 的变化说明在土壤轻旱(40%～45%)条件下,甘薯叶片气体交换主要受气孔因素限制,而重旱(30%～35%)主要受非气孔因素限制.施肥通过改善气孔因素对甘薯叶片气体交换产生促进效应.