Responses to water stress in two Eucalyptus globulus clones differing in drought tolerance

We evaluated drought resistance mechanisms in a drought-tolerant clone (CN5) and a drought-sensitive clone (ST51) of Eucalyptus globulus Labill. based on the responses to drought of some physiological, biophysical and morphological characteristics of container-grown plants, with particular emphasis on root growth and hydraulic properties. Water loss in excess of that supplied to the containers led to a general decrease in growth and significant reductions in leaf area ratio, specific leaf area and leaf-to-root area ratio. Root hydraulic conductance and leaf-specific hydraulic conductance decreased as water stress became more severe. During the experiment, the drought-resistant CN5 clone maintained higher leaf water status (higher predawn and midday leaf water potentials), sustained a higher growth rate (new leaf area expansion and root growth) and displayed greater carbon allocation to the root system and lower leaf-to-root area ratio than the drought-sensitive ST51 clone. Clone CN5 possessed higher stomatal conductances at moderate stress as well as higher hydraulic conductances than Clone ST51. Differences in the response to drought in root biomass, coupled with changes in hydraulic properties, accounted for the clonal differences in drought tolerance, allowing Clone CN5 to balance transpiration and water absorption during drought treatment and thereby prolong the period of active carbon assimilation.     

Physiological and biochemical responses to low non-freezing temperature of two Eucalyptus globulus clones differing in drought resistance

We have compared the metabolic responses of leaves and roots of two Eucalyptus globulus L. clones CN5 and ST51 that differ in their sensitivity to water deficits (ST51 is more drought sensitive), with regard to the effect of chilling (10/5 °C, day/night). We studied changes in growth, osmotic potential and osmotically active compounds, soluble proteins, leaf pigments, and membrane lipid composition. Our data showed that both clones have the ability to acclimatize to chilling temperatures. As a result of 10 days of acclimation, an increase of soluble sugars in leaves of treated plants of both clones was observed that disappeared later on. Differences between clones were observed in the photosynthetic pigments and soluble protein content which were more stable in CN5 under chilling. It also was apparent that CN5 presented a less negative predawn water potential (ψ_pd) and a higher leaf turgor than ST51 throughout the chilling treatment. In the case of the CN5, increased total lipids (TEA) and concomitant increase of linolenic acid (C18:3) in leaves after acclimatization may be related to a better clone performance under chilling temperatures. Moreover, a higher constitutive investment in roots in the case of CN5 as compared to ST51 may benefit new root regeneration under low temperatures favoring growth after cold Mediterranean winter.     

Growth, leaf morphology, water use and tissue water relations of Eucalyptus globulus clones in response to water deficit

Changes in leaf size, specific leaf area (SLA), transpiration and tissue water relations were studied in leaves of rooted cuttings of selected clones of Eucalyptus globulus Labill. subjected to well-watered or drought conditions in a greenhouse. Significant differences between clones were found in leaf expansion and transpiration. There was a significant clone x treatment interaction on SLA. Water stress significantly reduced osmotic potential at the turgor loss point (Pi0) and at full turgor (Pi100), and significantly increased relative water content at the turgor loss point and maximum bulk elastic modulus. Differences in tissue water relations between clones were significant only in the mild drought treatment. Among clones in the drought treatments, the highest leaf expansion and the highest increase in transpiration during the experiment were measured in those clones that showed an early and large decrease in Pi0 and Pi100.     

Leaf growth of Eucalyptus globulus seedlings under water deficit

Eucalyptus globulus Labill. seedlings grown under field conditions in Portugal were watered either daily (control) or every 6 days (drought-treated). Relative to those of control plants, rates of leaf production and leaf biomass accumulation were reduced by almost half in drought-treated plants. However, whereas expansion of new leaves on control plants slowed toward the end of the 30 day experiment, expansion of leaves of the same age on drought-treated plants accelerated as a change in weather conditions resulted in midday plant water potentials above -3.0 MPa. In plants that were left unwatered until they wilted and were then watered daily, expansion of the fifth leaf pair from the apex was slower than that of the same pair of leaves of plants watered daily throughout; but it continued for about twice as long and resulted in the same final leaf area. Drought treatment also caused a substantial reduction in the rate of leaf production, which, in part, accounted for the effect of drought on leaf biomass production. In a greenhouse study, witholding water for 15 days had only a slight effect on the length or width of adaxial epidermal cells, and the effect was quickly reversed on rewatering.     

Leaf water relations of Eucalyptus globulus ssp. globulus and E. nitens: seasonal, drought and species effects

In August 1990, a 2-ha plantation was established in an area where rainfall (about 515 mm year(-1)) was insufficient to meet evaporative demand. On nine occasions between September 1991 and April 1993, pressure-volume curves were constructed for irrigated and rainfed Eucalyptus globulus ssp. globulus Labill. and E. nitens (Deane and Maiden) Maiden trees. During the experiment, rainfed trees experienced six periods when predawn water potential was significantly lower than that of irrigated trees. In early spring of 1991 and 1992, osmotic potentials at full turgor and turgor loss point in the irrigated E. nitens were significantly lower than at other times of the year, probably because of winter hardening. Water stress reduced osmotic potential and increased bulk elastic modulus in E. nitens, whereas the reverse occurred in E. globulus. However, treatment differences with respect to changes in osmotic and elastic properties were commonly overshadowed by interspecific differences. These were most apparent at the end of the sixth period of water stress when osmotic potentials at full and zero turgor were significantly higher and bulk elastic modulus and relative water content at turgor loss point were significantly lower in E. globulus than in E. nitens. We conclude that the drought-tolerance responses of E. globulus make it a more suitable species than E. nitens for establishment on sites where moderate water stress is experienced.     

Growth and water relations of seedlings of two subspecies of Eucalyptus globulus

Seedlings of Eucalyptus globulus Labill subsp. globulus grown in soil in pots in the greenhouse grew faster than seedlings of E. globulus subsp. bicostata, and responded better to added nutrients and water. However, water stress caused a greater reduction in the growth of shoots and roots, and in the root/shoot ratios of fertilized seedlings of subsp. globulus than in those of bicostata. More leaf surface wax was produced by seedlings grown in the presence of fertilizer and an adequate supply of water than by seedlings subjected to nutrient or water stress. Despite larger amounts of leaf surface wax, seedlings of subsp. bicostata had higher epidermal conductances than seedlings of subspecies globulus. However, epidermal conductances were reduced more by water stress and by fertilization in seedlings of subsp. bicostata than in subsp. globulus. Tissue osmotic potentials at full and zero turgor were reduced by water stress only in seedlings of subsp. bicostata and were increased by fertilizer only in seedlings of subsp. globulus. The results indicate that although seedlings of subsp. globulus have inherently higher growth rates, seedlings of subsp. bicostata are better adapted to drought.     

Leaf water relations of Eucalyptus cloeziana and Eucalyptus argophloia in response to water deficit

Leaf water relations responses to limited water supply were determined in 7-month-old plants of a dry inland provenance of Eucalyptus argophloia Blakely and in a humid coastal provenance (Gympie) and a dry inland provenance (Hungry Hills) of Eucalyptus cloeziana F. Muell. Each provenance of E. cloeziana exhibited a lower relative water content at the turgor loss point, a lower apoplastic water content, a smaller ratio of dry mass to turgid mass and a lower bulk modulus of elasticity than the single provenance of E. argophloia. Osmotic potential at full turgor and water potential at the turgor loss point were significantly lower in E. argophloia and the inland provenance of E. cloeziana than in the coastal provenance of E. cloeziana. There was limited osmotic adjustment in response to soil drying in E. cloeziana, but not in E. argophloia. Between-species differences in water relations parameters were larger than those between the E. cloeziana provenances. Both E. cloeziana provenances maintained turgor under moderate water stress through a combination of osmotic and elastic adjustments. Eucalyptus argophloia had more rigid cell walls and reached lower water potentials with less reduction in relative water content than either of the E. cloeziana provenances, thereby enabling it to extract water from dryer soils.     

Ecophysiological and morphological responses to shade and drought in two contrasting ecotypes of Prunus serotina

Photosynthesis (A), water relations and stomatal reactivity during drought, and leaf morphology were evaluated on 2-year-old, sun- and shade-grown Prunus serotina Ehrh. seedlings of a mesic Pennsylvania seed source and a more xeric Wisconsin source. Wisconsin plants maintained higher A and leaf conductance (g(wv)) than Pennsylvania plants during the entire drought under sun conditions, and during the mid stages of drought under shade conditions. Compared to shade plants, sun plants of both sources exhibited a more rapid decrease in A or % A(max) with decreasing leaf water potential (Psi). Tissue water relations parameters were generally not significantly different between seed sources. However, osmotic potentials were lower in sun than shade plants under well-watered conditions. Following drought, shade plants, but not sun plants, exhibited significant osmotic adjustment. Sun leaves had greater thickness, specific mass, area and stomatal density and lower guard cell length than shade leaves in one or both sources. Wisconsin sun leaves were seemingly more xerophytic with greater thickness, specific mass, and guard cell length than Pennsylvania sun leaves. No source differences in leaf structure were exhibited in shade plants. Stomatal reactivity to sun-shade cycles was similar between ecotypes. However, well-watered and droughted plants differed in stomatal reactivity within and between multiple sun-shade cycles. The observed ecotypic and phenotypic variations in ecophysiology and morphology are consistent with the ability of Prunus serotina to survive in greatly contrasting environments.     

Applying PROMOD spatially across Tasmania with sensitivity analysis to screen for prospective Eucalyptus globulus plantation sites

When screening areas as prospective plantation sites two criteria are of interest: the expected yield at that site and the uncertainty associated with this yield expectation. This paper describes a methodology to enable spatial estimates of uncertainty to be attached to predictions of tree growth in relation to variations in soil depth, nutrient status, and drainage. An environmental sensitivity analysis was applied using the growth model PROMOD to produce site productivity and suitability maps for Eucalyptus globulus (Labill.) on a 1000 m regular grid of Tasmania. Nine productivity surfaces were computed, one for each of three assumed soil-water holding capacities (for soil depths of 0.5, 1.0, and 1.5 m — the common classes in Tasmania), combined with three permutations of soil-nutrient rating (the expected rating and one class above and one below this value), and incorporating potential waterlogging estimates. The mean and coefficient of variation was calculated for each 1000 m grid cell in the nine subsequent productivity surfaces. A final plantation suitability grid was then mapped to show areas of the State with predicted high (>25 m³ ha⁻¹per year), moderate (15–25 m³ ha⁻¹ per year), and low productivity (<15 m³ ha⁻¹ per year), each with corresponding high variability (coefficient of variation (CV>30%)) and low variability classes (CV<30%). As no areas had both, high productivity and high variability, only five plantation suitability classes were mapped.Those areas indicated as high productivity/low variation sites have a high probability of being successful E. globulus plantations. Low productivity sites with either high or low variation are very unlikely to be favourable for this species. Moderate productivity sites with low variation are likely to be viable potential plantation sites with relatively low risk, particularly with nutrient amelioration. Moderate productivity sites with high variation might warrant further field investigation to determine if there are major environmental factors or combinations of these, such as soil depth (water holding capacity), nutrient status, or drainage that are actually going to be limiting reasonable growth at these sites.     

Physiological and morphological responses to water stress in two Acacia species from contrasting habitats

Container-grown seedlings of Acacia tortilis Forsk. Hayne and A. xanthophloea Benth. were watered either every other day (well watered) or every 7 days (water-stressed) for 1 year in a greenhouse. Total plant dry mass (T(dm)), carbon allocation and water relations were measured monthly. Differences in leaf area (LA) accounted for differences in T(dm) between the species, and between well-watered and water-stressed plants. Reduction in LA as a result of water stress was attributed to reduced leaf initiation, leaf growth rate and leaf size. When subjected to prolonged water stress, Acacia xanthophloea wilted more rapidly than A. tortilis and, unlike A. tortilis, lost both leaves and branches. These differences between species were attributed to differences in the allocation of carbon between leaves and roots and in the ability to adjust osmotically. Rapid recovery in A. xanthophloea following the prolonged water-stress treatment was attributed to high cell wall elasticity. Previous exposure to water stress contributed to water-stress resistance and improved recovery after stress.     

Defence responses in plantation‐grown Eucalyptus globulus and Eucalyptus nitens after artificial fungal inoculation

The formation of reaction and barrier zones was studied in the xylem of Eucalyptus globulus and Eucalyptus nitens tree stems after wounding and artificial inoculation with two white rot fungi. The study had two objectives: to describe host responses in Eucalyptus spp. by light microscopy and to determine whether they would differ in a fungal treatment (wounding and inoculation by one of two fungal isolates) when compared to a control treatment (wounding only). Eucalyptus globulus and E. nitens developed similar reaction and barrier zones. The E. globulus barrier zone was characterized by kino vein formation. In both hosts, the reaction zone was primarily influenced by content and distribution of living tracheids and parenchyma cells within the sapwood. By contrast, the anatomy of the barrier zone showed similarities to the basic xylem structure of each host, except for some cell types that were newly formed (sclereids, kino veins) or increased in number (parenchyma cells, tracheids). Other cell types were reduced in number or completely absent. Host response in terms of barrier zone width appeared to be greater in the fungal than control treatment. Both wood decay fungi appeared to induce a wider barrier zone in both species than that associated with non‐specific damage caused exclusively by wounding. However, the small number of replicates available for this study was possibly insufficient to provide statistical evidence for different barrier zone width between fungal and control treatments.     

Differential physiological and morphological responses of two hybrid Populus clones to water stress

Hardwood cuttings of Populus clones Tristis #1 and Eugenei were grown in pots in a controlled environment chamber to observe early patterns of growth and water relations in response to changing conditions of water stress. Height and dry weight growth, dry matter partitioning, leaf area production, stomatal conductance and leaf water potential were measured periodically during the 73-day experiment. The two clones reacted in a similar way to an initial period of stress, showing reduced growth, stomatal conductance and leaf water potential. However, when stress was interrupted by ample watering and then reimposed, substantial differences between the clones were evident. Growth of Eugenei fully recovered after stress was relieved, especially leaf growth, but when water deficiency was reimposed, the plants wilted and some leaves died. Tristis #1, in contrast, showed a greater adaptation to changing stress conditions; it grew less than Eugenei after drought was interrupted, but showed little adverse effect of renewed water deficits. These responses were partially explained by the higher root/leaf weight ratios of Tristis #1 which enabled it to maintain a more favorable plant water status than Eugenei.     

Host responses to natural infection by Cytonaema sp. in the aerial bark of Eucalyptus globulus

The chemical and anatomical host responses to natural fungal infection by Cytonaema sp. in the aerial bark of 3‐year‐old Eucalyptus globulus plantation trees were examined. The lesion margin (LM) of the canker‐infected bark was characterized by the formation of a layer of dark extractives visible to the naked eye. Chemical analysis of the LM by gas chromatography‐mass spectrometry (GC‐MS), high‐performance liquid chromatography (HPLC) with ultraviolet detection and HPLC‐MS using negative ion electrospray ionisation indicated the presence of a range of compounds including hydrolysable tannins, polymeric proanthocyanidins, flavonoid glycosides, formlyated phloroglucinol compounds and volatile terpenes. These compounds were either undetectable in healthy tissue or present at significantly lower concentrations than in the LM. The LM of the canker‐infected bark was morphologically distinct from healthy phloem, its characteristics varying depending on severity of canker infection. In superficial infections in which only the phloem was affected, the following LMs were observed: (i) a continuous wound periderm of multiple layers, or (ii) an incompletely differentiated and discontinuous wound periderm. In cases of severe canker infections in which the vascular cambium had been killed, the new phloem formed subsequently contained traumatic oil glands in addition to the responses observed for superficial canker infections. All LMs were characterized by the formation of new parenchyma cells that stained positive for the presence of polyphenols. The significance of the chemical and structural responses as defence mechanisms against fungi causing stem canker is discussed.     

Hydraulic redistribution of soil water during summer drought in two contrasting Pacific Northwest coniferous forests

The magnitude of hydraulic redistribution of soil water by roots and its impact on soil water balance were estimated by monitoring time courses of soil water status at multiple depths and root sap flow under drought conditions in a dry ponderosa pine (Pinus ponderosa Dougl. ex Laws) ecosystem and in a moist Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) ecosystem. The fate of deuterated water applied to small plots to create a strong horizontal soil water potential gradient was also monitored to assess the potential for horizontal redistribution of water and utilization of redistributed water by co-occurring shallow-rooted plants. In a 20-year-old Douglas-fir stand, approximately 28% of the water removed daily from the upper 2 m of soil was replaced by nocturnal hydraulic redistribution during late August. In an old-growth ponderosa pine stand, approximately 35% of the total daily water utilization from the upper 2 m of soil appeared to be replaced by hydraulic redistribution during July and August. By late September, hydraulic redistribution in the ponderosa pine stand was no longer apparent, even though total water use from the upper 2 m of soil was nearly identical to that observed earlier. Based on these results, hydraulic redistribution would allow 21 and 16 additional days of stored water to remain in the upper soil horizons in the ponderosa pine and Douglas-fir stands, respectively, after a 60-day drought. At both sites, localized applications of deuterated water induced strong reversal of root sap flow and caused soil water content to cease declining or even temporarily increase at locations too distant from the site of water application to have been influenced by movement of water through the soil without facilitation by roots. Xylem water deuterium values of ponderosa pine seedlings suggested utilization of redistributed water. Therefore, hydraulic redistribution may enhance seedling survival and maintain overstory transpiration during summer drought. These first approximations of the extent of hydraulic redistribution in these ecosystems suggest that it is likely to be an important process in both wet and dry forests of the Pacific Northwest.     

Ecophysiological responses of a young blue gum (Eucalyptus globulus) plantation to weed control

Early weed control may improve the growth of forest plantations by influencing soil water and nutrient availability. To understand eucalypt growth responses to weed control, we examined the temporal responses of leaf gas-exchange, leaf nitrogen concentration (N) and water status of 7-month-old Eucalyptus globulus L. trees in a paired-plot field trial. In addition, we monitored the growth, leaf N and water status of the competing vegetation in the weed treatment. By the end of the 11-month experiment, complete weed control (WF treatment) of largely woody competitors increased the basal diameter of E. globulus by 14%. As indicated by pre-dawn water potentials of >?-?0.05 MPa, interspecies competition for water resources was minimal at this site. In contrast, competition for N appeared to be the major factor limiting growth. Estimations of total plot leaf N (g m(-2) ground) showed that competing vegetation accounted for up to 70% of the total leaf N at the start of the trial. This value fell to 15% by the end of the trial. Despite increased leaf N(area) in WF trees 5 months after imposition of weed control, the photosynthetic capacity (A(1500)) of E. globulus was unaffected by treatment suggesting that the growth gains from weed control were largely unrelated to changes in leaf-level photosynthesis. Increased nutrient availability brought about by weed control enabled trees to increase investment into leaf-area production. Estimates of whole-tree carbon budget based on direct measurements of dark respiration and A(1500) allowed us to clearly demonstrate the importance of leaf area driving greater productivity following early weed control in a nutrient-limited site.     

Anatomical and histochemical defence responses induced in juvenile leaves of Eucalyptus globulus and Eucalyptus nitens by Mycosphaerella infection

The responses of juvenile leaves of two Eucalyptus species, with contrasting susceptibility to infection by Mycosphaerella leaf disease, were compared. The anatomical changes, accumulation of phenolics, suberin, lignin and anthocyanin and the retention of chlorophyll were studied in leaf lesions of varying developmental stages caused by species of Mycosphaerella. Enhanced resistance of Eucalyptus nitens in southern Australia was attributed to the formation of an effective lignified and suberized necrophylactic periderm, to restrict pathogen spread. Leaves of E. nitens contained isobilateral palisade which resulted in both abaxial and adaxial cell division and the initiation of a strong reinforced cellular zone from an early lesion stage. Eucalyptus globulus formed a slower, distorted necrophylactic periderm through hypertrophic changes to existing mesophyll and limited cell divisions of the single adaxial palisade layer. Deposits of lignin and suberin did not occur until later in lesion development, which were not effective in preventing further disease development. From this study it is hypothesized that tolerance of eucalypts to Mycosphaerella pathogens may be associated with constitutive mesophyll density.     

Carbon isotope discrimination and its relationship to drought resistance under field conditions in genotypes of Eucalyptus globulus Labill.

The aim of this study was to investigate the relationship between carbon isotope discrimination (Δ), survival and growth, to determine the value of Δ for screening for yield improvement under drought conditions. Intra-specific variation in carbon isotope discrimination (Δ) was examined in Eucalyptus globulus. Wood samples were collected from 3 to 7 year-old populations from three different field-sites at Huelva (south-western Spain). Differences between study sites were related to the availability of water resulting from soil physical properties. Average carbon isotope discrimination ranged from 16.7 to 18.1‰. There was a 2.2‰ maximum range in Δ values among different trees of the same age. A strong, positive correlation was observed between average carbon isotope discrimination and average tree diameter at breast height (DBH_ob). Average carbon isotope discrimination was also strongly correlated with survival at the Valdeoscuro site (r²=0.95) and with the percentage of failures in neighbouring trees (r²=0.65) at the San Sebastián site. These results suggest a strong effect of availability of water on Δ values. Implications for the use of carbon isotope discrimination in selecting for better adapted genotypes are discussed.     

Assessing genetic variation to improve stem straightness in Eucalyptus globulus

Context

Stem straightness is an important trait for growers and processors of Eucalyptus globulus logs for solid-wood products.

Aims

The aims of the study were to determine the extent of genetic variation in stem deviation from straightness in E. globulus and assess the utility of a six-point subjective scoring method as a selection criterion for stem straightness.

Methods

Two E. globulus progeny trials, grown under solid-wood product regimes, were studied. At age 9 years (post-thinning), stem straightness was measured using both image analysis and a six-point subjective scale. Diameter at breast height (DBH; 1.3 m) was measured at both age 5 (pre-thinning) and age 9 years.

Results

Significant additive genetic variation was observed. Strong, positive and significant additive genetic correlations were observed between the stem straightness assessment methods and between DBH at ages 5 and 9 years. Significant positive genetic correlations were shown between subjectively scored stem straightness and DBH at both ages 5 and 9 years.

Conclusion

The six-point subjective scoring method is a cost-effective selection criterion for stem straightness in E. globulus. The image measurement technique may be applied where objective estimates of stem straightness are required, for training purposes and to verify subjective scores.     

Association of Eucalyptus globulus leaf anatomy with susceptibility to Teratosphaeria leaf disease

Resistance of Eucalyptus globulus juvenile foliage to Teratosphaeria leaf disease (TLD) damage has been shown to be under genetic control. Constitutive traits of juvenile leaves such as stomatal density (counted with wax on and with wax removed), total phenolics, total leaf wax, leaf mass per area (LMA) and leaf anatomical features were assessed to determine differences between pairs of resistant and susceptible families. Anatomic features assessed included cuticle, epidermis, palisade spongy mesophyll and total leaf thickness, as well as palisade cell size, shape and intercellular airspace. One interprovenance, one intraprovenance and two within‐family contrasts were used to compare resistant and susceptible pairs of families of Australian origin. The more resistant families had a significantly higher LMA and smaller percentage airspace compared with susceptible families. It is argued that one of the mechanisms by which E. globulus resists TLD involves the closely packed palisade mesophyll cells that increases LMA and reduces airspace, thereby restricting pathogen spread once infection has occurred.     

Mycosphaerella species associated with leaf disease of Eucalyptus globulus in Ethiopia

Eucalyptus spp. are among the most widely planted exotic trees in Ethiopia. Several damaging leaf pathogens are known from Eucalyptus spp. worldwide. Of these, Mycosphaerella spp. are among the most important, causing the disease known as Mycosphaerella leaf disease (MLD). Characteristic symptoms of MLD include leaf spot, premature defoliation, shoot and twig dieback. Recent disease surveys conducted in Ethiopian Eucalyptus plantations have revealed disease symptoms similar to those caused by Mycosphaerella spp. These symptoms were restricted to E. globulus trees growing in several localities in south, south western and western Ethiopia. The aim of this study was to identify the fungi associated with this disease. This was achieved by examining ascospore germination patterns, anamorph associations and sequence data from the Internal Transcribed Spacer (ITS) region of the rRNA operon, for representative isolates. Several different ascospore germination patterns were observed, suggesting that more than one species of Mycosphaerella is responsible for MLD on E. globulus in Ethiopia. Analysis of sequence data showed that three Mycosphaerella spp., M. marksii, M. nubilosa and M. parva were present. This is the first report of these three species from Ethiopia and represents a valuable basis on which to build further studies in the region.