Different vulnerabilities of Quercus ilex L. to freeze- and summer drought-induced xylem embolism: an ecological interpretation

Quercus ilex L. growing in the southern Mediterranean Basin region is exposed to xylem embolism induced by both winter freezing and summer drought. The distribution of the species in Sicily could be explained in terms of the different vulnerability to embolism of its xylem conduits. Naturally occurring climatic conditions were simulated by: (1) maintaining plants for 3h at ambient temperatures of 0, -1.5, -2.5, -5.0 and -11°C; and (2) allowing plants to dry out to ratios of their minimum diurnal leaf water potentials (Ψ₁) to that at the turgor loss point (Ψ_tlp) of 0.6, 0.9, 1.05, 1.20 and 1.33. The loss of hydraulic conductivity of one-year-old twigs reached 40% at -1.5°C and at Ψ₁/Ψ_tlP= 1.05. Recovery from these strains was almost complete 24 h after the release of thermal stress or after one irrigation, respectively. More severe stresses reduced recovery consistently. The percentages of xylem conduits embolized following application of the two stresses, were positively related to xylem conduit diameter. The capability of the xylem conduits to recover from stress was positively related to the conduit diameter in plants subjected to summer drought, but not in the plants subjected to winter freezing stress. The ecological significance of the different vulnerabilities to embolism of xylem conduits under naturally occurring climatic conditions is discussed.     

Hydraulic architecture of leaf venation in Laurus nobilis L.

Veins are the main irrigation system of the leaf lamina and an understanding of the hydraulic architecture of the vein networks is essential for understanding leaf function. However, determination of leaf hydraulic parameters is challenging, because for most leaves the vein system is reticulate, contains a hierarchy of different vein sizes, and consists of leaky conduits. We present a new approach that allows for measurements of pressure differences between the petiole and any vein within the leaf. Measurements of Laurus nobilis leaves indicate that first‐ and second‐order veins have high axial conductance and relatively small radial permeability, thus allowing water to reach distal areas of the leaf with only a small loss of water potential. Higher order veins tend to be more hydraulically resistant and permit greater radial leakage. This design allows for a relatively equitable distribution of water potential and thus reflects the capacity of the venation to provide a relatively homogeneous water supply across the leaf lamina, with only the leaf margins being hydraulically disadvantaged relative to the rest of the leaf.     

Phylogeographical variation of chloroplast DNA in holm oak (Quercus ilex L.)

Variation in the lengths of restriction fragments (RFLPs) of the whole chloroplast DNA molecule was studied in 174 populations of Quercus ilex L. sampled over the entire distribution of this evergreen and mainly Mediterranean oak species. By using five endonucleases, 323 distinct fragments were obtained. From the 29 and 17 cpDNA changes identified as site and length mutations, respectively, 25 distinct chlorotypes were distinguished, mapped and treated cladistically with a parsimony analysis, using as an outgroup Q. alnifolia Poech, a closely related evergreen oak species endemic to Cyprus where Q. ilex does not grow. The predominant role of Q. ilex as maternal parent in hybridization with other species was reflected by the occurrence of a single very specific lineage of related chlorotypes, the most ancestral and recent ones being located in the southeastern and in the northwestern parts of the species’ geographical distribution, respectively. The lineage was constituted of two clusters of chlorotypes observed in the ‘ilex’ morphotyped populations of the Balkan and Italian Peninsulas (including the contiguous French Riviera), respectively. A third cluster was divided into two subclusters identified in the ‘rotundifolia’ morphotyped populations of North Africa, and of Iberia and the adjacent French regions, respectively. Postglacial colonization probably started from three distinct southerly refugia located in each of the three European peninsulas, and a contact area between the Italian and the Iberian migration routes was identified in the Rhône valley (France). Chlorotypes identical or related to those of the Iberian cluster were identified in the populations from Catalonia and the French Languedoc region, which showed intermediate morphotypes, and in the French Atlantic populations which possessed the ‘ilex’ morphotype, suggesting the occurrence of adaptive morphological changes in the northern part of the species’ distribution.     

Leaf temperature effects on gas exchange in Quercus ilex L. growing under field conditions

ABSTRACT

Gas exchange temperature dependence in Quercus ilex shrubs growing in the Mediterranean maquis was analysed. The gas exchange trend was monitored during the year: photosynthetic activity (A _net) reached the highest average rates in early spring and autumn (12.5 µmol m^-2s^-1 was the absolute maximum A _net measured) and the lowest rates were monitored in the middle of June. There was a good correlation (r = 0.72) between A _net and g _s (A _net = 4.1246 ln g _s + 4316; P < 0.01), indicating that stomatal control of CO₂ diffusion plays an important role in controlling photosynthetic activity. Leaf temperature allowing the highest photosynthetic and stomatal conductance rates of Quercus ilex were in the range 17.5 – 29°C. A _net and g_s dropped below half its maximum value when leaf temperatures were below 11.5°C and above 35.7°C. Transpiration rates (E) were strongly related to leaf temperature; E increased as leaf temperature increased and the highest E rates were monitored in June, despite a 46% decrease in g _s. Leaf water loss from transpiration, during the drought period, could result in leaf water stress which would exacerbate heat effects on photosynthesis. During summer, the increase in leaf temperatures decreased g _s which in turn decreased A _net. Consequently, stomatal control in Quercus ilex may be considered as an adaptive strategy during drought.     

New evidence for a role of vessel-associated cells and phloem in the rapid xylem refilling of cavitated stems of Laurus nobilis L.

Xylem recovery from embolism was studied in Laurus nobilis L. stems that were induced to cavitate by combining negative xylem pressure potentials (P_X = ?1.1 MPa) with positive air pressures (P_C) applied using a pressure collar. Xylem refilling was measured by recording the percentage loss of hydraulic conductance (PLC) with respect to the maximum 2 min, 20 min and 15 h after pressure release. Sodium orthovanadate (an inhibitor of many ATP‐ases) strongly inhibited xylem refilling while fusicoccin (a stimulator of the plasma membrane H⁺‐ATPase) promoted complete embolism reversal. So, the refilling process was interpreted to result from energy‐dependent mechanisms. Stem girdling induced progressively larger inhibition to refilling the nearer to the embolized stem segment phloem was removed. The starch content of wood parenchyma was estimated as percentages of ray and vasicentric cells with high starch content with respect to the total, before and after stem embolism was induced. A closely linear positive relationship was found to exist between recovery from PLC and starch hydrolysis. This, was especially evident in vasicentric cells. A mechanism for xylem refilling based upon starch to sugar conversion and transport into embolized conduits, assisted by phloem pressure‐driven radial mass flow is proposed.     

Purification and Characterization of a Magnesium Ion Requiring N-Acetyl-D-glucosamine Specific Lectin from Seeds of Quercus ilex L.

A new magnesium ion requiring N-acetyl-D-glucosamine specific lectin QIL was purified to electrophoretic homogeneity from seeds of Quercus ilex L. through successive steps of (i) lectin extraction, (ii) ammonium sulphate (30–50%) fractionation, (iii) diethylaminoethyl (DEAE)-cellulose chromatography, (iv) carboxymethyl (CM)-cellulose chromatography, and (v) Sephadex G-75 chromatography. The lectin, having specific activity of 25,600 hemagglutination units (HAU)/mg of protein, was found to be a monomeric protein with a native molecular weight of 13.2 kDa. N-Acetyl-D-glucosamine was found to exhibit most potent inhibitory action on the lectin activity among all the sugars tested. The lectin was also found to exhibit specificity for human blood groups A, B, and AB. It was converted to the corresponding apo-lectin by ethylenediaminetetraacetic acid (EDTA) treatment followed by buffer dialysis. The apo-lectin exhibited a specific and characteristic requirement for magnesium ions for the expression of its activity.     

夏休闲期复种油菜对旱地土壤水分和小麦产量的影响

以冬小麦收获后高留茬休闲地为对照,连续4年在陇东黄土旱塬设6个油菜播种期,研究了休闲期作物覆盖对土壤水分及后作冬小麦产量与水分利用效率的影响.结果表明：不同播种期夏休闲期土壤贮水量差异显著(P＜0.05),其中8月5日播种油菜的土壤蓄水效率为58.5%,产量和水分利用效率较对照提高7.5%和5.9%.平均而言,夏休闲期复种油菜后作小麦干旱年份增产16.1%,平水年份增产6.8%.夏休闲期复种油菜是西北旱地小麦抗旱增产的有益途径.     

不同栽培模式对冬小麦-夏玉米轮作系统产量、氮素累积和平衡的影响

以河北山前平原区秸秆还田条件下小麦-玉米轮作体系为研究对象,设置农民习惯、高产高效、再高产和再高产高效4个模式,通过定位试验探讨各栽培模式对3个轮作周期作物产量、土壤硝态氮累积量及氮平衡的影响.结果表明: 小麦、玉米产量均以再高产模式最高,高产高效和再高产高效模式次之,均显著高于农民习惯模式;小麦季和玉米季氮肥利用效率(PFP)均以高产高效模式最高,显著高于其他模式;0～400 cm土体硝态氮累积量在 768.4～1133.3 kg·hm^-2之间,其中80%～85%累积在根下90～400 cm土层;4种模式的土壤硝态氮均有明显向下淋移现象,120～150 cm和270～330 cm处均出现了累积峰,以270～330 cm土层硝态氮累积量最大;高产高效模式的土壤硝态氮含量整体水平均低于其他模式,浓度基本维持在30 mg·kg^-1以下,在一定程度上能有效缓解环境压力;冬小麦季0～90 cm土体氮素盈余量均小于夏玉米季,并以高产高效模式的氮素表观损失量最低,显著低于其他模式.综合考虑产量、氮肥利用效率、硝态氮累积和氮平衡,以高产高效模式表现最优,但还有一定的提升空间.     

Variation in onset of summer dormancy and flowering capacity along an aridity gradient in Poa bulbosa L., a geophytic perennial grass

Variation in the onset of summer dormancy and flowering capacity of 16 populations of Poa bulbosa, collected along a steep north-south aridity gradient in Israel (810-110 mm rain year(-1)), was studied under controlled conditions in a phytotron (16 h daylength, 22/16 degrees C day/night) and under natural conditions in a garden experiment in a net-house. Plant age at the onset of dormancy varied markedly amongst populations (7-16 weeks under controlled conditions) and was positively correlated with mean annual precipitation at the site of origin of the population, i.e. dormancy was earlier as aridity increased. Flowering capacity in the different populations was negatively correlated with rainfall in the original habitat and, consequently, also with the age at onset of dormancy, i.e. the lower the mean annual precipitation, the earlier the onset of dormancy and the higher the proportion of flowering plants and panicles per plant. Differences in xeromorphic leaf traits were also observed among populations from locations differing in aridity. Plants from the more arid sites (110-310 mm year(-1)) generally had greyish and curved leaves, whereas plants from more humid sites (500-810 mm year(-1)) tended to have green and straight leaves. Thus, plants with curved and/or greyish leaves generally had a higher flowering capacity and entered dormancy earlier than plants with straight and/or green leaves. The significance of the association among these traits for the adaptation of P. bulbosa to increasing aridity is discussed.     

Long-term exposure to elevated [CO2] in a natural Quercus ilex L. community: net photosynthesis and photochemical efficiency of PSII at different levels of water stress

Naturally grown trees of Mediterranean evergreen oak (Quercus ilex L.), representing the climax species of the region, were enclosed in six large open-top chambers and exposed to ambient and elevated CO₂ concentrations during a 3 year period. Maximum daily net photosynthetic rates measured at the two different CO₂ concentrations were from 30 to 100% higher in elevated than in ambient [CO₂] throughout the experimental period. The increase in maximum daily photosynthesis was also accompanied by a 93% rise in the apparent quantum yield of CO₂ assimilation, measured during periods of optimum soil moisture conditions. Hence, no clear evidence of down-regulation of net photosynthetic activity was found. Interactions between atmospheric CO₂ concentration and plant water stress were studied by following the natural evolution of drought in different seasons and years. At each level of water stress, the maximum rate of carbon assimilation was higher in elevated than in ambient [CO₂] by up to 100%. Analysis of in vivo chlorophyll fluorescence parameters in normal (21%) and low (2%) oxygen concentrations provided useful insights into the functioning and stability of the photosynthetic processes. The photochemical efficiency of PSII (F_v/F_m) progressively decreased as drought conditions became more evident; this trend was accentuated under elevated [CO₂]. Thermal de-excitation processes were possibly more significant under elevated than under ambient [CO₂], in a combination of environmental stresses. This research suggests two possible conclusions: (i) a ‘positive’ interaction between elevated [CO₂] and carbon metabolism can be obtained through relief of water stress limitation in the summer months, and (ii) elevated [CO₂], under drought conditions, may also enhance the significance of slow-relaxing quenching.     

Photo- and antioxidative protection during summer leaf senescence in Pistacia lentiscus L. grown under Mediterranean field conditions

Summer leaf senescence in Pistacia lentiscus L. plants serves to remobilize nutrients from the oldest leaves to the youngest ones, and therefore contributes to plant survival during the adverse climatic conditions typical of Mediterranean summers, i.e. water deficit superimposed on high solar radiation and high temperatures. To evaluate the extent of photo- and antioxidative protection during leaf senescence of this species, changes in carotenoids, including xanthophyll cycle pigments, and in the levels of ascorbate and alpha-tocopherol were measured prior to and during summer leaf senescence in 3-year-old plants grown under Mediterranean field conditions. Although a chlorophyll loss of approx. 20% was observed during the first stages of leaf senescence, no damage to the photosynthetic apparatus occurred as indicated by constant maximum efficiencies of photosystem II photochemistry. During this period the de-epoxidation state of the xanthophyll cycle, and lutein, neoxanthin and ascorbate levels were kept constant. At the same time beta-carotene and alpha-tocopherol levels increased by approx. 9 and 70%, respectively, presumably conferring photo- and antioxidative protection to the photosynthetic apparatus. By contrast, during the later stages of leaf senescence, characterized by severe chlorophyll loss, carotenoids were moderately degraded (neoxanthin by approx. 20%, and both lutein and beta-carotene by approx. 35%), ascorbate decreased by approx. 80% and alpha-tocopherol was not detected in senescing leaves. This study demonstrates that mechanisms of photo- and antioxidative protection may play a major role in maintaining chloroplast function during the first stages of leaf senescence, while antioxidant defences are lost during the latest stages of senescence.     

麦季牛场肥水灌溉对冬小麦-夏玉米产量与磷吸收利用及土壤剖面分布的影响

在华北平原灌溉区,采用冬小麦 夏玉米周年轮作田间试验,研究麦季牛场肥水灌溉对冬小麦和夏玉米产量、磷吸收量、磷累计利用率及土壤积累的影响.结果表明：麦季肥水灌溉能显著提高冬小麦和夏玉米产量,冬小麦产量随肥水带入磷的增加先增加后降低,肥水灌溉带入137 kg P₂O₅·hm^-2时冬小麦产量最高,磷的当季利用率较高,分别为7646.4 kg·hm^-2和24.8%,肥水灌溉带入过量磷会降低冬小麦产量和磷当季利用率;夏玉米产量和磷素吸收量随冬小麦季肥水灌溉带入磷量增加而增加,后季夏玉米产量增加2222.4～2628.6 kg·hm^-2,磷吸收量增加13.9～21.1 kg·hm^-2.农民习惯施肥处理夏玉米当季施磷88 kg P₂O₅·hm^-2时,与不施肥处理相比,夏玉米产量增加2235.0 kg·hm^-2.随着牛场肥水灌溉年限的推移,作物增产效果逐渐明显,冬小麦 夏玉米轮作体系作物累计磷利用率逐年升高,6季作物收获后,磷累计利用率达40.0%～47.7%.试验条件下,
冬小麦 夏玉米轮作体系进行2次肥水灌溉是较经济安全的灌溉模式.     

干旱胁迫对菜苔叶片保护酶活性和膜脂过氧化的影响

以3个不同耐旱性的菜苔（Brassica parachinensis L.H.Bailey）品种为试材,研究了干旱胁迫对叶片保护酶活性和膜脂过氧化的影响及其与抗旱性的关系。干旱胁迫条件下,菜苔叶片的电解质外渗率和MDA含量呈上升趋势,叶绿素含量、抗坏血酸含量和SOD活性呈下降趋势,CAT活性表现为先上升后下降。耐旱品种比不耐旱品种具有较高的叶绿素含量和抗坏血酸含量,具有较低的电解质外渗率和MDA含量;耐旱品种的SOD活性比不耐旱品种下降幅度小。轻度干旱胁迫下,耐旱品种的CAT活性上升幅度比不耐旱品种高;重度干旱胁迫下耐旱品种的CAT活性下降程度比不耐旱品种低。耐旱品种的POD活性在干旱条件下先上升而后降低,不耐品种的POD活性处于下降趋势。干旱6d后,耐旱品种的SOD、CAT和POD活性显著高于不耐旱品种。     

长期浅耕与秸秆还田对关中平原冬小麦-夏玉米轮作土壤钾素含量及层化比率的影响

采用土壤分层采样和钾素化学形态分组方法,探讨连续13年秸秆全量还田与浅耕对关中平原冬小麦 夏玉米轮作体系土壤各形态钾素含量及层化比率的影响. 结果表明： 采样区土壤速效钾和非交换性钾均有显著的表聚现象和层化效应,速效钾及其组分表现尤为突出,且15～30 cm土层亏缺严重,已低于缺钾临界值;速效钾及其组分的土壤表层(0～5 cm)与5～15 cm和15～30 cm耕层土壤的比值(SR₁与SR₂)之间呈显著性差异,非交换性钾与矿物钾的比值则差异不显著. 连续秸秆全量还田对维持冬小麦 夏玉米轮作体系土壤钾库平衡有积极贡献,但应高度关注有效钾素表聚和层化对土壤钾肥力的负面效应.     

Survival and recovery of perennial forage grasses under prolonged Mediterranean drought: I. Growth, death, water relations and solute content in herbage and stubble

Swards of Dactylis glomerata cultivars (cvs) KM2 and Lutetia and of Lolium perenne cvs Aurora and Vigor were grown under full irrigation or prolonged summer drought (80 d) in a field experiment in the South of France.
After irrigation was withheld, leaf extension rates of all cvs fell by 90% within 9–12 d, and rapid scorching of laminae followed. Tiller mortality at the end of the drought was very different in the cocksfoot cvs (4% for KM2 and 76% for Lutetia) and intermediate (41%) for both ryegrass cvs. Following re-watering, rates of herbage regrowth were closely correlated with tiller survival. Measured minerals contributed c . 0·52 MPa to osmotic potential in all treatments, whereas water-soluble carbohydrates (WSC) contributed 0·25 MPa under irrigation and 0·46 MPa during drought.
There was no systematic difference between the two species for summer survival under severe drought, but large differences between the cocksfoot cvs. The traits most strongly associated with superior survival were: (a) a deep root system and greater water uptake at depth; (b) low water and osmotic potentials in surviving laminae, i.e. better tolerance to dehydration; (c) large pool-size of WSC reserves (fructans having degree of polymerization >4) in entire tiller bases (stubble); (d) low accumulation of proline in stubble; (e) rapid nitrogen uptake after rewatering.     

Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought

A comparative study on stomatal control under water deficit was conducted on grapevines of the cultivars Grenache, of Mediterranean origin, and Syrah of mesic origin, grown near Montpellier, France and Geisenheim, Germany. Syrah maintained similar maximum stomatal conductance (g_max) and maximum leaf photosynthesis (A_max) values than Grenache at lower predawn leaf water potentials, Ψ_leaf, throughout the season. The Ψ_leaf of Syrah decreased strongly during the day and was lower in stressed than in watered plants, showing anisohydric stomatal behaviour. In contrast, Grenache showed isohydric stomatal behaviour in which Ψ_leaf did not drop significantly below the minimum Ψ_leaf of watered plants. When g was plotted versus leaf specific hydraulic conductance, K_l, incorporating leaf transpiration rate and whole‐plant water potential gradients, previous differences between varieties disappeared both on a seasonal and diurnal scale. This suggested that isohydric and anisohydric behaviour could be regulated by hydraulic conductance. Pressure‐flow measurements on excised organs from plants not previously stressed revealed that Grenache had a two‐ to three‐fold larger hydraulic conductance per unit path length (K_h) and a four‐ to six‐fold larger leaf area specific conductivity (LSC) in leaf petioles than Syrah. Differences between internodes were only apparent for LSC and were much smaller. Cavitation detected as ultrasound acoustic emissions on air‐dried shoots showed higher rates for Grenache than Syrah during the early phases of the dry‐down. It is hypothesized that the differences in water‐conducting capacity of stems and especially petioles may be at the origin of the near‐isohydric and anisohydric behaviour of g.