Coexistent Mediterranean woody species as a driving factor of Phytophthora cinnamomi infectivity and survival

The long-term conservation of Mediterranean mixed oak forests is seriously threatened by the massive mortality of Quercus suber caused by the exotic pathogen Phytophthora cinnamomi. This species frequently grows in mixed forests under natural conditions, but nothing is known about how its level of disease might be altered by the diversity and identity of coexisting neighbours varying in susceptibility to the exotic pathogen. Here we analysed the individual and combined effects of Q. suber and the main coexisting tree species (Quercus canariensis and Olea europaea subsp. europaea var. sylvestris) in mixed forests of southern Spain on the production of infective and survival spores of P. cinnamomi. Through in vitro experiments, it was demonstrated that mixtures of Q. suber and Q. canariensis highly stimulated the production of P. cinnamomi zoospores in comparison with both species in monocultures. Olea europaea did not stimulate zoospore production. Under controlled conditions, the initial and final densities of inoculum in soil planted with monocultures of O. europaea and Q. canariensis did not differ. However, inoculum densities significantly decreased along the experiment in Q. suber mixtures with O. europaea and Q. canariensis. Phytophthora cinnamomi was able to infect and cause root rot symptoms on all tree species, including O. europaea var. sylvestris. We concluded that mixed stands of Q. suber and Q. canariensis are able to stimulate P. cinnamomi infectivity and survival much more than monospecific stands, and consequently under favourable conditions for root disease development, the coexistence of Q. suber and Q. canariensis might exacerbate Mediterranean forests decline. This study also constitutes the first report of O. europaea var. sylvestris as host and inductor of P. cinnamomi sporulation under controlled conditions.     

Trunk injection of fosetyl‐aluminium controls the root disease caused by Phytophthora cinnamomi on Quercus ilex woodlands

In Spain, Quercus open woodlands are animal ranching systems of organic production seriously threatened by the exotic pathogen Phytophthora cinnamomi. The root disease it causes kills thousands of oaks annually. Effective disease management needs to integrate different techniques, and the use of a resistance inducer such as fosetyl‐Al can play a key role, because the use of potassium phosphite is prohibited in Spain. In a woodland where the pathogen recently arrived, 60 holm oaks in three different defoliation classes (asymptomatic, slight and moderate defoliation) were selected for trunk injection with pressurised capsules containing 4% of commercial fosetyl‐Al or water (controls). Holm oaks were checked periodically for defoliation and presence of the pathogen in roots and rhizosphere soil. Three years after treatments, defoliation was significantly lower in oaks treated with fosetyl‐Al, which even increased canopy cover, in comparison with control oaks, independent of the initial defoliation class considered. Chlamydospore density in rhizosphere soil, as well as the presence of the pathogen into the roots, was not significantly influenced by fosetyl‐Al treatments, although a trend to a lower presence of P. cinnamomi in roots was observed in treated oaks at every soil inoculum density detected. This study has shown that fosetyl‐Al, a phosphonate registered as a fungicide in the European Union, provides protection to holm oaks against P. cinnamomi, even exhibiting a therapeutic effect on pre‐existing infections. Consequently, this effective measure should be considered as part of the integrated approach to control this highly destructive pathogen in holm oak woodlands.     

Long‐term directional changes in upland Quercus forests throughout Oklahoma,USA

Questions: (1) How have the composition and structure of undisturbed upland Quercus forests changed over 50 years across a large region and moisture gradient; (2) What factors are associated with long‐term and broad‐scale changes in these forests? Location: Oklahoma, USA. Methods: We re‐sampled 30 forest stands originally sampled in the 1950s across a large geographical area and compared basal area, tree density, and sapling density between the sampling periods using paired t‐tests, CCA, and DCA. We examined vegetation dynamics in the context of drought indices compiled for the sample period. Results: Total and Quercus stellata basal area and tree density increased, but Q. stellata and Q. marilandica sapling density decreased. Juniperus virginiana and woody species richness increased for all measures. DCA indicated that re‐sampled stands generally changed from Q. stellata–Q. marilandica‐dominated forests to forests with greater woody species richness and more J. virginiana. Q. stellata remained a dominant tree species; otherwise, composition shifted towards mesophytic and invasive woody species. Measurements taken in the 1950s immediately followed a major drought; whereas subsequent decades were significantly moister. Conclusions: Fire exclusion and drought may have played an important role in driving changes towards lower dominance by Quercus, increased importance of mesophytic and invasive species, and greater woody species richness. These phenomena are similar to those found in Quercus‐dominated forests throughout the northern hemisphere.     

Combined effects of soil properties and Phytophthora cinnamomi infections on Quercus ilex decline

Aims

The importance of soil properties as determinants of tree vitality and Phytophthora cinnamomi root infections was analysed.

Methods

The study comprised 96 declining stands in western Spain, where declining and non-declining holm oak (Quercus ilex L.) trees were sampled. Soil properties (soil depth, Ah horizon thickness, texture, pH, redox potential, soil bulk density and N-NH₄ ⁺ and N-NO₃ ^? concentrations) and P. cinnamomi infections were assessed.

Results

Tree mortality rates increased with low soil bulk densities, which were also associated with more P. cinnamomi-infected trees. Occurrence of infected trees was higher in fine textured soils and in thick Ah horizons. Fine textured soils favoured trees, but with the presence of P. cinnamomi their health status deteriorated. Soil under declining trees had higher N-NO₃ ^?/N-NH₄ ⁺ ratio values than under non-declining trees. Additional soil properties changes associated to grazing were not related to decline and P. cinnamomi infections.

Conclusions

The implications of P. cinnamomi in holm oak decline and the influence of soil properties as contributors to pathogen activity were demonstrated. Fine soil textures and thick Ah horizons, usually favourable for vigour and vitality of trees growing in the Mediterranean climate, were shown to be disadvantageous soil properties if P. cinnamomi was present. Fine soil textures and thick Ah horizons are frequently related with higher levels of soil moisture, which increase the inoculum of the pathogen and favours root infection. Grazing does not seem to be directly linked to Q. ilex health status or P. cinnamomi root rot.     

The Balearic Islands: a reservoir of cpDNA genetic variation for evergreen oaks

Aim To analyse the role of the Balearic Islands as a refuge area for evergreen Quercus (cork oak: Quercus suber L., holm oak: Q. ilex L., kermes oak: Q. coccifera L.), by using molecular, historical and palaeobotanical data. Location The Western Mediterranean Basin (Balearic Islands, eastern Iberia, Provence, Sardinia, Corsica, Sicily, Malta, Italy, Northern Africa). Methods We sampled 108 populations and used the PCR‐RFLP technique with five universal cpDNA primers to define haplotypes in the sampled populations. Diversity, differentiation parameters and spatial analysis of the populations, using a spatial version of amova , were linked to the geological history of the Western Mediterranean Basin in order to explain the present spatial pattern of the evergreen Quercus populations in the Balearics. Results Evergreen Quercus cpDNA shows a complex structure, with remnants of ancient diversity in the Balearics. Balearic populations of holm oak are related to Iberian populations, while for cork and kermes oaks, we found both Tyrrhenian and Iberian haplotypes. Main conclusions The complex spatial patterns of cpDNA in Balearic evergreen Quercus appears explicable in terms of a combination of physical (vicariance and long distance dispersal) and biological (introgressive hybridization) factors. The Balearics constitute a glacial refuge area and a reservoir of genetic variation with traces of ancient diversity from Messinian–Pliocene stages.     

Defining Plant Resistance to Phytophthora cinnamomi: A Standardized Approach to Assessment

Phytophthora cinnamomi is a soil‐borne plant pathogen that causes devastating disease in agricultural and natural systems worldwide. While a small number of species survive infection by the pathogen without producing disease symptoms, the nature of resistance, especially under controlled conditions, remains poorly understood. At present, there are no standardized criteria by which resistance or susceptibility to P. cinnamomi can be assessed, and we have used five parameters consisting of plant fresh weight, root growth, lesion length, relative chlorophyll content of leaves and pathogen colonization of roots to analyse responses to the pathogen. The parameters were tested using two plant species, Zea mays and Lupinus angustifolius, through a time course study of the interactions and resistance and susceptibility defined 7 days after inoculation. A scoring system was devised to enable differentiation of these responses. In the resistant interaction with Z. mays, there was no significant difference in fresh weight, root length and relative chlorophyll content in inoculated compared with control plants. Both lesion size and pathogen colonization of root tissues were limited to the site of inoculation. Following inoculation L. angustifolius showed a significant reduction in plant fresh weight and relative leaf chlorophyll content, cessation of root growth and increased lesion lengths and pathogen colonization. We propose that this technique provides a standardized method for plant–P. cinnamomi interactions that could be widely used to differentiate resistant from susceptible species.     

Regulation of defence responses in avocado roots infected with Phytophthora cinnamomi (Rands)

Phytophthora cinnamomi occurs worldwide and has a host range in excess of 1,000 plant species. Avocados (Persea americana Mill) have been described as highly susceptible to this soil-borne pathogen. Here, the regulation of defence responses in avocado root seedlings inoculated with P. cinnamomi mycelia is described. A burst of reactive oxygen species (ROS) was observed 4 days after inoculation. The higher physiological concentration of H₂O₂ induced by P. cinnamomi on avocado roots had no effect on in vitro growth of the oomycete. Total phenols and epicathecin content showed a significant decrease, but lignin and pyocianidins exhibited no changes after inoculation. Also, increased nitric oxide (NO) production was observed 72 h after treatment. We studied the effects of one NO donor [sodium nitroprusside (SNP)], and one NO scavenger [2- to 4-carboxyphenyl-4,4,5,5-tetramethylimidazole-1-oxyl-3-oxide (CPTIO)] to determine the role of NO during root colonisation by P. cinnamomi mycelia. Pretreatment of the roots with CPTIO, but not with SNP, inhibited root colonisation suggesting an important role for NO production during the avocado–P. cinnamomi interaction. Our data suggest that although defence responses are activated in avocado roots in response to P. cinnamomi infection, these are not sufficient to avoid pathogen invasion.     

Effect of Brassica Biofumigant Amendments on Different Stages of the Life Cycle of Phytophthora cinnamomi

The oomycete plant pathogen Phytophthora cinnamomi causes a highly destructive root rot that affects numerous hosts. Integrated management strategies are needed to control P. cinnamomi in seminatural oak rangelands. We tested how biofumigation affects crucial stages of the pathogen's life cycle in vitro, in infested soils under laboratory conditions and in planta. Different genotypes of three potential biofumigant plant species (Brassica carinata, Brassica juncea, Brassica napus) were collected at different phenological stages, analysed for their glucosinolate contents, and subsequently tested. The most effective genotypes against mycelial growth and sporangial production were further tested on the viability of chlamydospores in artificially infested natural soils and in planta on Lupinus luteus, a host highly susceptible to P.cinnamomi. Brassica carinata and B. juncea genotypes inhibited mycelial growth, decreased sporangial production, and effectively inhibited the viability of chlamydospores in soil, but only B. carinata decreased disease symptoms in plants. Effective genotypes of Brassica had high levels of the glucosinolate sinigrin. Biofumigation with Brassica plants rich in sinigrin has potential to be a suitable tool for control of oak root disease caused by P. cinnamomi in Spanish oak rangeland ecosystems.     

Type designation of the oaks Quercus ballota and Q. rotundifolia (Fagaceae)

The names Quercus ballota and Q. rotundifolia (Fagaceae), referring to taxa sometimes considered conspecific with the widely distributed Q. ilex, are discussed and typified. A specimen preserved at MPU is designated as the lectotype of Desfontaines's name Q. ballota. Lamarck's name Q. rotundifolia is neotypified using a specimen preserved at VAL, with several duplicates in other European herbaria.     

湖南壳斗科栎属一新种——德夯栎

该文报道了采自湖南省湘西世界地质公园德夯园区壳斗科栎属一新种——德夯栎(Quercus dehangensis G. X. Chen, D. G. Zhang & B. Z. Wang),根据其壳斗小苞片覆瓦状排列而被归于栎属。德夯栎和巴东栎(Quercus engleriana Seem.)相近,但与巴东栎的区别在于前者植株通常3~5 m高,侧脉7~10对,小枝、叶柄及叶背被覆具鳞片的星状毛,托叶早落。根据IUCN红色名录标准,德夯栎濒危等级为极危(CR)。     

The Holocene and Upper Pleistocene pollen sequence of Carihuela Cave, southern Spain

A new pollen sequence (ca. 15,700-1250 yr BP) is presented for three stratigraphical sections of Carihuela Cave (Granada, southeastern Spain), thus completing a record that covers from the last Interglacial to late Holocene. The Late Glacial is characterized by open landscapes with junipers and early colonisation of Quercus, while the Holocene is depicted by mixed oak forests, with a diversity of broad-leaf trees and scrub, which decrease after ca. 5470 yr BP synchronously with the expansion of xerophytes and occurrence of indicators of anthropogenic disturbance. The whole pollen record of Carihuela fits into the general trends described for reference pollen sites of southern Europe, including Padul in the province of Granada, and other sequences from Mediterranean Spain, through which the heterogeneity of environmental change increases from mid to late Holocene. We conclude that, in contrast with other regions of Spain, deciduous Quercus-dominated forests are very old in eastern Andalusia, thus conflicting with floristic phytosociological models of vegetation change that imply that monospecific Q. ilex/rotundifolia woodlands are the potential mature forest in the region. Dating results suggest that the last Neanderthals of Carihuela lived between ca. 28,440 and 21,430 yr BP, which agrees with the postulation that southern Spain was the latest refugium for this human species in Europe.     

Root Foraging Influences Plant Growth Responses to Earthworm Foraging

Interactions among the foraging behaviours of co-occurring animal species can impact population and community dynamics; the consequences of interactions between plant and animal foraging behaviours have received less attention. In North American forests, invasions by European earthworms have led to substantial changes in plant community composition. Changes in leaf litter have been identified as a critical indirect mechanism driving earthworm impacts on plants. However, there has been limited examination of the direct effects of earthworm burrowing on plant growth. Here we show a novel second pathway exists, whereby earthworms (Lumbricus terrestris L.) impact plant root foraging. In a mini-rhizotron experiment, roots occurred more frequently in burrows and soil cracks than in the soil matrix. The roots of Achillea millefolium L. preferentially occupied earthworm burrows, where nutrient availability was presumably higher than in cracks due to earthworm excreta. In contrast, the roots of Campanula rotundifolia L. were less likely to occur in burrows. This shift in root behaviour was associated with a 30% decline in the overall biomass of C. rotundifolia when earthworms were present. Our results indicate earthworm impacts on plant foraging can occur indirectly via physical and chemical changes to the soil and directly via root consumption or abrasion and thus may be one factor influencing plant growth and community change following earthworm invasion. More generally, this work demonstrates the potential for interactions to occur between the foraging behaviours of plants and soil animals and emphasizes the importance of integrating behavioural understanding in foraging studies involving plants.     

The Microscopic Examination of Phytophthora cinnamomi in Plant Tissues Using Fluorescent In Situ Hybridization

The microscopic examination of Phytophthora cinnamomi in plant tissues is often difficult as structures such as hyphae, chlamydospores and oospores are frequently indistinguishable from those of other fungi and oomycetes, with histological stains not enabling species differentiation. This lack of staining specificity makes the localization of P. cinnamomi hyphae and reproductive structures within plant tissues difficult, especially in woody tissues. This study demonstrates that with the use of a species‐specific fluorescently labelled DNA probe, P. cinnamomi can be specifically detected and visualized directly using fluorescent in situ hybridization (FISH) without damage to plant or pathogen cell integrity or the need for subculturing. This approach provides a new application for FISH with potential use in the detailed study of plant–pathogen interactions in plants.     

Stomatal and Mesophyll Limitations to Photosynthesis in One Evergreen and One Deciduous Mediterranean Oak Species

In the evergreen Quercus rotundifolia and the co-existing deciduous Q. faginea we studied the diurnal variations in photosynthetic capacity (P _max), measured as the rate of O₂ evolution at photon and CO₂ saturation, and in the rate of net CO₂ assimilation (P _N) in the field during the period of maximum photosynthetic activity. Our aim was to check the contribution of stomatal and non-stomatal limitations to the diurnal variation in photosynthesis, and to study the differences between both species. Q. faginea leaves displayed lower mass per unit area and higher nitrogen content than Q. rotundifolia leaves. The maximum stomatal conductance and P _N in the field were higher in Q. faginea than in Q rotundifolia. Also P _max of Q. faginea was higher than that of Q. rotundifolia. Both species attained in the field a high percentage of the P _max (around 82 % for Q. faginea and 73 % for Q. rotundifolia). This indicates reduced stomatal limitation of photosynthesis under favourable conditions, especially in Q. faginea. P _N underwent a sharp decrease towards mid-day in association with increase in the atmospheric vapour pressure deficit and decrease in the leaf water potential. P _max was also reduced during mid-day. This demonstrated the contribution of mesophyll limitations to the P _N in the two species under stress. The mesophyll limitation of photosynthesis seemed to be similar for both species, independently from the differences in leaf traits between them.     

Genetic diversity in oak populations under intensive management for fuelwood in the Sierra de Zongolica,Mexico

Firewood and charcoal are used on a daily basis both in rural areas and in cities. This type of energy is produced by one of the most ancient traditional methods, known as coppice, which harvest tree sprouts. There is controversy about its effects on forests: it preserves populations and tree cover of species used, but reduces density, inhibits sexual reproduction and generates genetic erosion. We inquired if it was possible to identify a loss of genetic diversity in oak populations traditionally used for charcoal by the Zongolica Nahuas in Veracruz state, Mexico. We studied populations of Quercus laurina, Quercus calophylla and Quercus rugosa in three different altitudes. Molecular analysis with eight nuclear codominant microsatellites was performed to determine the diversity, structure and gene flow of these species. Results for Q. laurina were Na = 8.458, I = 1.766, Ho = 0.679, polymorphism = 100%, Fis = 0.079, with intraindividual variation of 81.55%. For Q. calophylla: Na = 7.250, I = 1.563, Ho = 0.646, polymorphism = 91.67%, Fis = 0.083, with intraindividual variation of 83.80%. For Q. rugosa: Na = 6.958, I = 1.510, Ho = 0.574, polymorphism = 91.67%, Fis = 0.204, with intraindividual variation of 81.99%; this species shows signals of an early genetic isolation process. Our findings indicate that Quercus genetic diversity for the three species is high and comparable with oak species in Mexico and worldwide. We conclude that at the present, coppice is preserving a historical diversity in adult trees kept alive through sprouting. Nonetheless, problems with coppice systems elsewhere, unregulated harvesting and expansion of pine plantation in the region suggest that further studies, hand in hand with a landscape management approach that improve charcoal and firewood production, may be valuable for Sierra de Zongolica genetic biodiversity conservation.     

Sexual reproduction in the cork oak (Quercus suber L). II. Crossing intra- and interspecific barriers

Intraspecific barriers promote outcrossing while interspecific mechanisms may contribute to the isolation of species; both control the exchange of genes between plants. In this paper we establish that post-pollination mechanisms promote outcross and act at different temporal and spatial levels to control seed set and quality in Quercus species. Controlled pollinations were performed to investigate intraspecific crossing barriers in Q. suber and pollen-pistil interactions following interspecific pollinations with some simpatric Quercus species. Cytological data of intraspecific crosses in Q. suber and Q. ilex have shown different kinetics on pollen tube growth after self and outcross pollination, but pollen tubes were able to reach the base of the styles in both species and seed set was successful. Although some pre-zygotic interaction is occurring at the style, the most important interaction takes place at ovary. The cross between Q. ilex and Q. suber is possible only in one direction, revealing a case of unilateral incongruity. We show that the lack of seed set observed in the cross Q. suber×Q. ilex is due to the inability of pollen tubes to penetrate the transmitting tissue after germination. With Q. suber mainly as female parent, pollinations with other simpatric Quercus species have shown different levels of constraint on pollen tube progression at stigma and style levels. Eventual hybridisation between Quercus species will depend on the compatibility of pollen-pistil interactions, on the competitive ability of pollen genotypes, and, most important, on the overlapping of geographical, phenological and ecological factors. Differences in seed set and seed allocation was evident in all crosses, and was particularly outstanding in interspecific and in self intraspecific crosses, determining the ultimate level of seed production and quality in Quercus species. Received: 4 April 2001 / Accepted: 8 May 2001     

辽东栎幼苗根系形态特征对环境梯度的响应

以辽东栎(Quercus liaotungensis)在陕西不同分布区:秦岭北坡(太白)、黄土高原南部(黄龙)和黄土高原中部(延安)为研究地点并设置样地,对1—5年生辽东栎幼苗的根系形态指标进行测定,分析辽东栎幼苗根系形态特征及其与环境因子的关系。结果表明:由秦岭北坡到黄土高原中部,黄龙地区辽东栎幼苗根系在发育前期(1—2年)低于太白和延安,总体上黄龙地区幼苗根系总长度、表面积、总体积、根尖数、平均直径、组织密度和单株生物量高于太白和延安地区。在太白地区,辽东栎幼苗根系表面积、总体积和平均直径较小,根系分岔数较大,幼苗根系主要通过提高分岔数来拓展自己的营养空间以适应环境;在黄龙和延安地区,幼苗根系表面积、总体积和平均直径较大,根系分岔数较小,幼苗根系主要是通过根系的伸长生长适应胁迫环境。3个地区辽东栎幼苗根系总长度、表面积、总体积、根尖数和分岔数随年龄的增长呈线性函数变化格局,均可用线性函数方程y=ax+b(a0,P0.05)进行描述。冗余分析表明幼苗根系分岔数、总长度、比根长和根尖数与土壤速效磷、硝态氮、速效钾、降雨量、石砾含量和速效氮呈正相关;与较高的土壤pH值、年均温和夏季气温呈负相关。未来辽东栎林抚育经营中,含石砾的湿润土壤生境更有利于辽东栎幼苗根系生长。     

Differential ecophysiological responses to seasonal drought of three co-existing oak species in northern Greece

Oaks (Quercus spp.) represent the most important broadleaf genus with respect to forest-shaping tree species in the Mediterranean. Considering future climate scenarios (increased drought conditions), the identification of drought tolerant oak species is of great importance for future forest management in this region. The objective of the study was the comparison of physiological status of three economically and ecologically valuable oak species (Quercus ilex, Quercus frainetto and Quercus pubescens) co-existing in natural coppice stands in NE Greece, in response to seasonal drought stress. Measurements were conducted between June and September 2016, every 15–20 days until leaf falling. The parameters studied were predawn leaf water potential and fast chlorophyll fluorescence induction curves (OJIP test), chlorophyll content, and relative water content. Meteorological data from the area were also collected. Photosynthetic parameters such as performance indices (PI_abs and PI_tot) reacted to summer drought conditions, with Q. frainetto showing the lowest values. The discrepancy between species increased with duration of drought period. Q. frainetto revealed the lowest predawn water potential values. The results indicate that Q. frainetto is less suitable for future forestry applications in the studied climate/elevation zone than Q. pubescens and Q. ilex.     

A comparison of leaf physiology and anatomy of Quercus (section Erythrobalanus-Fagaceae) species in different light environments

Physiological and anatomical attributes of leaves were examined of three species of Quercus section Erythrobalanus. All three species occur in moist temperate deciduous forests of eastern North America. Seedlings of each species were grown in different light conditions for comparison. The attributes measured were net photosynthesis, stomatal conductivity, blade and cuticle thickness, stomatal density, thickness of upper and lower epidermis, and thickness of palisade mesophyll. The results generally demonstrate the close association between anatomical adaptations and efficiency of physiological processes; they also elucidate the distribution patterns of the three Quercus species across the forest topography in southern New England. The most drought-tolerant and light-demanding species, Q. velutina (Lam.), exhibited the greatest leaf anatomical plasticity, the highest net photosynthesis in the different light conditions, and the lowest stomatal area per unit area of leaf. The most drought-intolerant species, Q. rubra (L.), exhibited the least leaf anatomical plasticity, the lowest net photosynthesis in the different light conditions, and the highest stomatal area per unit area of leaf. Quercus coccinea (Muenchh.) usually exhibited values that were intermediate between Q. rubra and Q. velutina.