Elemental composition and flavonol content of <Emphasis Type="Italic">Pentaphylloides fruticosa</Emphasis> (L.) O. Schwarz in connection with plant age

The elemental composition and flavonol content of Pentaphylloides fruticosa (L.) O. Schwartz under the conditions of the Altai mountains are investigated. Some differences in the concentrations of elements and flavonols were detected in the leaves of P. fruticosa plants of different age classes corresponding to two periods of development: pregenerative (immature and virginal plants) and generative (young, middle-aged and old generative individuals). The maximum content of flavonols, in particular quercetin, in the leaves of P. fruticosa corresponds to the young generative age. A very high correlation between the flavonol content and the concentrations of Mn, Zn, Mo, and Se was revealed.     

Leaf age affects the quality of DNA extracted from Dimorphandra mollis (Fabaceae), a tropical tree species from the Cerrado region of Brazil

Isolation of high-quality DNA from plants, especially plants from the Cerrado, is notoriously difficult because of polysaccharides and secondary compounds produced by plants from this biome. DNA isolation and its quality may be compromised by chemical defenses such as tannins and phenols. Quantitative plant defenses tend to have a cumulative effect, increasing in concentration during leaf development, reducing DNA quality extracted in mature compared to young leaves. We report the effect of leaf age on DNA extraction of Dimorphandra mollis. Our working hypothesis was that the young leaves have more DNA than old leaves of the same individual because chemical defenses accumulate in older leaves. Young and old leaves were sampled from eight mature trees as well as leaves from eight seedlings in the north region of Minas Gerais State. Genomic DNA extraction followed the standard CTAB procedure. DNA isolation was very successful from young leaves of 16 individuals of D. mollis. The extracted DNA exhibited high quality and the DNA quantity was also high, with an A(260)/A(280) ratio above 1.8, which is within the optimal sample range. In contrast, DNA isolation from old leaves was not successful. When the DNA was extracted from old leaves, the DNA was brownish, indicating contamination by phenolic compounds. These metabolites oxidize the DNA irreversibly, which hinders amplification of DNA by PCR by inhibiting the action of enzymes such as Taq polymerase. PCR performed with DNA from young leaves of D. mollis was successful and produced strong bands for RAPD markers.     

Seasonal variation in leaf characteristics and food selectionby larval noctuids on an evergreen Mediterranean shrub

Despite year round availability of foliage, abundance of generalist noctuid larvae (Lepidoptera: Noctuidae) in evergreen-dominated Mediterranean forests has a narrow, distinct spring peak. This restricted larval period has been suggested to result in part from avoidance of the nutritionally poor mature foliage, and preference for nutritionally superior spring-produced young leaves. This study examines this hypothesis by (i) documenting differences in nutritional characteristics between expanding (April) and mature (June) young leaves of the evergreen Mediterranean shrub Daphne laureola L. (Thymelaeaceae), and (ii) experimentally studying the feeding preferences of noctuid larvae for young leaves, old leaves (≥ 1 yr old), and developing fruits of this species in one south-eastern Spanish locality. Young leaves of D. laureola declined in nutrient concentration and specific dry mass from April to June. The responses of noctuid larvae, in terms of both relative preference and total consumption, to this seasonal variation in chemical and physical features of young leaves were also investigated. When noctuid larvae were simultaneously offered young leaves, old leaves and developing fruits, they exhibited similar preferences for young leaves and developing fruits, and rejected old leaves developed during the previous year. Noctuid larvae did not modify their consumption of young leaves relative to old leaves and developing fruits in response to seasonal changes. Food selection patterns exhibited by D. laureola noctuid herbivores, notably the rejection of old leaves in favour of young ones, are consistent with the hypothesis relating restricted larval periods of these generalist consumers with the low food value of the previous season leaves of evergreen Mediterranean plants.     

Phosphorus retranslocation in Hordeum vulgare during early tillering

Summary In Hordeum vulgare, phosphorus retranslocation was studied after it had been supplied to the roots for three days (experiment 1), and after foliar application (experiments 3–8). Phosphorus uptake by leaves of different ages was also measured 16 and 60 minutes after ³²P addition to the medium (experiment 2).In experiment 1, treatments at 0.6 and 31 p.p.m. of phosphorus were applied when the first leaf had completed its rapid growth. The plants were then grown for three days in media labelled with ³²P, and for a subsequent 10 days in non-labelled solutions. Retranslocation was measured by changes in total phosphorus and in ³²P.Both root feeding, and foliar application of ³²P, demonstrated three phases during leaf development: import (recently initiated leaf), export (mature leaf) and an intermediate phase with both export and import (leaf half developed).There was large transport of foliar applied ³²P, from mature leaves to roots, and some of this ³²P was re-exported to the shoots, including the mature leaves. Root feeding of ³²P over short periods strongly suggested that phosphorus uptake by the shoots occurred via the xylem, even at low phosphorus.In experiment 1, there were distinct treatment differences in relative growth rates, growth of young organs and roots, and in phosphorus concentrations of all but the very young leaves. Mature leaves showed a large net phosphorus export at low phosphorus, but a large net import at high phosphorus. This was not due to treatment differences in export, because total export from the mature leaves was even somewhat smaller at low than at high phosphorus. The treatment differences, with net export at low but net import at high phosphorus, were thus due to the higher import in the mature leaves at high phosphorus. Total export remained at a high level throughout the experiment at high phosphorus, while it declined with time at low phosphorus.For phosphorus absorbed during early growth, both the export from the mature leaves, and the intake by the developing leaves, was independent of phosphorus treatment; i.e. for each individual organ the quantities of phosphorus involved were the same in the two phosphorus treatments. Thus, the higher phosphorus contents of developing organs at high phosphorus were obtained from phosphorus supplied to the roots during later growth, and not from phosphorus supplied during early growth of the whole plant.The data are consistent with the notion that phosphorus export is controlled in the source. It is suggested that at high phosphorus this control is due to a saturation of the sites transporting phosphorus into the phloem. At low phosphorus, on the other hand, release from individual leaf cells might have been the dominating factor.     

Involvement of the thylakoidal NADH-plastoquinone-oxidoreductase complex in the early responses to ozone exposure of barley (Hordeum vulgare L.) seedlings

A possible implication of the plastid NADH-plastoquinone-oxidoreductase (Ndh) complex in the response against ozone-mediated oxidative stress in barley (Hordeum vulgare L.) leaves was investigated. After a 4 h treatment, exposure of barley seedlings to moderate ozone concentrations produced leaf-age-dependent increases in lipid peroxidation, peroxidase, and Ndh complex activities in the thylakoid membranes. A significant amount and activity of the Ndh complex were detected in mature barley leaves, but not in young barley leaves. In fact, young barley leaves behaved like ndh-deficient leaves in most of the aspects studied. When plants were exposed to photo-oxidative light after ozone fumigation, the recovery of Fv/Fm was lower in young leaves than in mature leaves. Ozone treatment significantly decreased non-photochemical quenching (qN) in young leaves, but not in mature leaves. Mature leaves showed higher levels of the energy (DeltamuH+) dependent (qE) component of qN. Treatment with antimycin A, an inhibitor of cyclic electron flow, increased the decay of qN produced by ozone in young leaves, but not in mature ones. The reduction state of plastoquinone increased after ozone treatment in mature dark-adapted leaves and was strongly quenched by far red light. It is proposed that the function of the Ndh complex helps the maintenance of qN, probably through the poising of the redox steady-state level of the intersystem carriers and then by optimizing the rate of cyclic electron flow. This should constitute an age-dependent early response in barley leaves, by contributing to minimize photoinhibition in the presence of ozone and high light.     

Determination of critical K concentrations in sap from individual leaves or from whole plants using K-interruption experiments

Summary A new method is described for estimating critical K concentrations from K interruption experiments using only 2 treatments. Frequent measurements are made of the growth and K concentration of plants subjected to either continued or interrupted K supply and the data used to define the relation between relative yield and K concentration for the K-deficient plants. Critical concentrations are estimated from the results using a mathematical model of plant growth to interpolate over the critical concentration region of the curve. The method has the advantage that the critical concentrations are determined at the exact time that growth is affected. The method was tested using data from previously published experiments with lettuce in which the concentrations of K were measured in sap from both the total shoot and from individual leaf petioles. The model accurately predicted the form of the relationship between relative yield and K concentration for the total shoot and for young expanding leaves, but consistently deviated from the data for recently matured ones. Average estimates of critical concentration ranged fromca. 18 to 34 mmoll⁻¹ in the young leaves and from 48 to 67 mmoll⁻¹ in the mature ones when Na salts were present or absent respectively. The values for total shoot sap were similar to those for mature leaves. The critical concentrations for young expanding leaves were virtually identical to the minimum believed to be needed for the maintenance of important biochemical processes in individual cells, and suggests that a single critical K concentration for plant sap might apply to a wide range of crops provided an actively growing part of the plant is sampled.     

Xylogalacturonan exists in cell walls from various tissues of Arabidopsis thaliana

Evidence is presented for the presence of xylogalacturonan (XGA) in Arabidopsis thaliana. This evidence was obtained by extraction of pectin from the seeds, root, stem, young leaves and mature leaves of A. thaliana, followed by treatment of these pectin extracts with xylogalacturonan hydrolase (XGH). Upon enzymatic treatment, XGA oligosaccharides were primarily produced from pectin extracts obtained from the young and mature leaves and to a lesser extent from those originating from the stem of A. thaliana. The oligosaccharide GalA(3)Xyl was predominantly formed from these pectin extracts. No XGA oligosaccharides were detected in digests of pectin extracts from the seeds and roots. A low number of XGA oligosaccharides was obtained from pectins of A. thaliana. This indicates a uniform distribution of xylose in XGA from A. thaliana. The predominant production of GalA(3)Xyl, as well as the release of linear GalA oligosaccharides pointed to a lower degree of xylose substitution in XGA from A. thaliana than in XGA from apple and potato. The estimated amount of XGA accounted for approximately 2.5%, 7% and 6% (w/w) of the total carbohydrate in the pectin fraction of the stem, young leaves and mature leaves, respectively.     

Photosynthetic regulation under fluctuating light in field-grown Cerasus cerasoides: A comparison of young and mature leaves

Photosystem I (PSI) is a potential target of photoinhibition under fluctuating light. However, photosynthetic regulation under fluctuating light in field-grown plants is little known. Furthermore, it is unclear how young leaves protect PSI against fluctuating light under natural field conditions. In the present study, we examined chlorophyll fluorescence, P700 redox state and the electrochromic shift signal in the young and mature leaves of field-grown Cerasus cerasoides (Rosaceae). Within the first seconds after any increase in light intensity, young leaves showed higher proton gradient (ΔpH) across the thylakoid membranes than the mature leaves, preventing over-reduction of PSI in the young leaves. As a result, PSI was more tolerant to fluctuating light in the young leaves than in the mature leaves. Interestingly, after transition from low to high light, the activity of cyclic electron flow (CEF) in young leaves increased first to a high level and then decreased to a stable value, while this rapid stimulation of CEF was not observed in the mature leaves. Furthermore, the over-reduction of PSI significantly stimulated CEF in the young leaves but not in the mature leaves. Taken together, within the first seconds after any increase in illumination, the stimulation of CEF favors the rapid lumen acidification and optimizes the PSI redox state in the young leaves, protecting PSI against photoinhibition under fluctuating light in field-grown plants.     

Leaf Photosynthesis,Dark Respiration and Fluorescence as Influenced by Leaf Age in an Evergreen Tree,Prosopis Juliflora

P. juliflora trees produce leaves during two growth periods. The first cohort of leaves is produced during spring in cool conditions, while the second cohort is produced during monsoon under warm conditions. I studied photosynthetic characteristics of young, mature, and old leaves of the previous season (monsoon) in the spring season. Maximum net photosynthetic rate of a young leaf was lower than that of the mature and old leaves. The total CO₂ fixed per day by the young leaves was just 36 % of that in the mature leaves while the old leaves fixed 76 % of that of the mature leaf. The total transpiration rate and water use efficiency (WUE) were similar in the mature and old leaves, while they were much lower in the young leaves. Dark respiration rate was maximal in the young leaves as compared to the mature and old leaves. About 92 % of the total CO₂ fixed per day were respired by the young leaves. The diurnal fluorescence characteristics (F/F_m, q _p, and q _N) of the young, mature, and old leaves showed that photochemical efficiency of photosystem 2 during midday decreased more in the young and old leaves than in the mature ones. However, the fluorescence characteristics showed that in all the three leaf types there was complete recovery of the photochemical efficiency at sunset from the midday depression. F_v/F_m in the young and mature leaves also confirmed this. Hence the young and old leaves were photosynthetically less efficient than mature leaves, but they were well adapted to withstand the harsh environmental conditions.     

Mechanisms for tolerance of very high tissue phosphorus concentrations in Ptilotus polystachyus

Study of plants with unusual phosphorus (P) physiology may assist development of more P‐efficient crops. Ptilotus polystachyus grows well at high P supply, when shoot P concentrations ( [P] ) may exceed 40 mg P g^?1 dry matter (DM). We explored the P physiology of P. polystachyus seedlings grown in nutrient solution with 0–5 mM P. In addition, young leaves and roots of soil‐grown plants were used for cryo‐scanning electron microscopy and X‐ray microanalysis. No P‐toxicity symptoms were observed, even at 5 mM P in solution. Shoot DM was similar at 0.1 and 1.0 mM P in solution, but was ～14% lower at 2 and 5 mM P. At 1 mM P, [P] was 36, 18, 14 and 11 mg P g^?1 DM in mature leaves, young leaves, stems and roots, respectively. Leaf potassium, calcium and magnesium concentrations increased with increasing P supply. Leaf epidermal and palisade mesophyll cells had similar [P]. The root epidermis and most cortical cells had senesced, even in young roots. We conclude that preferential accumulation of P in mature leaves, accumulation of balancing cations and uniform distribution of P across leaf cell types allow P. polystachyus to tolerate very high leaf [P].     

Photosynthetic Acclimation in Pea and Soybean to High Atmospheric CO2 Partial Pressure

Nonnodulated pea (Pisum sativum L. cv Frosty) and soybean (Glycine max [L.] Merr. cv Wye) plants were grown under artificial lights from germination with ample nutrients, 600 [mu]mol photons m-2 s-1, and either 34 to 36 (control) or 64 to 68 Pa (enriched) CO2. For soybean, pod removal and whole-plant shading treatments were used to alter the source-sink balance and carbohydrate status of the plants. Growth of both species was substantially increased by CO2 enrichment despite some down-regulation of photosynthesis rate per unit leaf area ("acclimation"). Acclimation was observed in young pea leaves but not old and in old soybean leaves but not young. Acclimation was neither evident in quantum yield nor was it related to triose phosphate limitation of net photosynthesis. A correlation between levels of starch and sugars in the leaf and the amount of acclimation was apparent but was loose and only weakly related to the source-sink balance of the plant. A consistent feature of acclimation was reduced ribulose bisphosphate carboxylase (RuBPCase) content, although in vivo RuBPCase activity was not necessarily diminished by elevated growth CO2 owing to increased percentage of activation of the enzyme. A proposal is discussed that the complexity of photosynthetic acclimation responses to elevated CO2 is as an expression of re-optimization of deployment of within-plant resources at three levels of competition.     

Systemic regulation of leaf anatomical structure, photosynthetic performance, and high-light tolerance in sorghum

Leaf anatomy of C3 plants is mainly regulated by a systemic irradiance signal. Since the anatomical features of C4 plants are different from that of C3 plants, we investigated whether the systemic irradiance signal regulates leaf anatomical structure and photosynthetic performance in sorghum (Sorghum bicolor), a C4 plant. Compared with growth under ambient conditions (A), no significant changes in anatomical structure were observed in newly developed leaves by shading young leaves alone (YS). Shading mature leaves (MS) or whole plants (S), on the other hand, caused shade-leaf anatomy in newly developed leaves. By contrast, chloroplast ultrastructure in developing leaves depended only on their local light conditions. Functionally, shading young leaves alone had little effect on their net photosynthetic capacity and stomatal conductance, but shading mature leaves or whole plants significantly decreased these two parameters in newly developed leaves. Specifically, the net photosynthetic rate in newly developed leaves exhibited a positive linear correlation with that of mature leaves, as did stomatal conductance. In MS and S treatments, newly developed leaves exhibited severe photoinhibition under high light. By contrast, newly developed leaves in A and YS treatments were more resistant to high light relative to those in MS- and S-treated seedlings. We suggest that (1) leaf anatomical structure, photosynthetic capacity, and high-light tolerance in newly developed sorghum leaves were regulated by a systemic irradiance signal from mature leaves; and (2) chloroplast ultrastructure only weakly influenced the development of photosynthetic capacity and high-light tolerance. The potential significance of the regulation by a systemic irradiance signal is discussed.     

Developmental changes in cell wall structure of phloem fibres of the bamboo Dendrocalamus asper

BACKGROUND AND AIMS: Bamboo culms have excellent physical and mechanical properties, which mainly depend on their fibre content and anatomical structure. One of the features which is known to contribute to the high tensile strength in bamboo is the multilayered structure of the fibre cell wall. The aim of this study was to characterize the development of the layered structure in fibre cell walls of developing and maturing culms of Dendrocalamus asper. METHODS: Cell wall development patterns were investigated in phloem fibre caps of vascular bundles in the inner culm wall areas of Dendrocalamus asper of three different age classes (<6 months old, 1 year old, 3 years old). A combination of light microscopy and image analysis techniques were employed to measure cell wall thickness and to determine number of cell wall layers, as well as to describe the layering structure of fibre walls. Two-dimensional maps showing the distribution pattern of fibres according to the number of cell wall layers were produced. KEY RESULTS: The cell walls of fibres in phloem fibre caps located in the inner part of the culm wall of D. asper developed rapidly during the first year of growth. Six different fibre types could be distinguished based upon their cell wall layering and all were already present in the young, 1-year-old culm. In the mature stage (3 years of age) the multilayering was independent of the cell wall thickness and even the thinner-walled fibres could have a large number of wall layers. The multilayered nature of cell wall structure varied considerably between individual cells and was not exclusively related to the cell wall thickness. Nevertheless, fibres at the periphery of the fibre bundles and immediately adjacent to the phloem elements exhibited a consistent and high degree of layering in their cell walls. CONCLUSIONS: The multilayered structure of fibre cell walls was formed mainly during the first year of growth by the deposition of new wall layers of variable thickness, resulting in a high degree of heterogeneity in the layering patterns amongst individual fibres. A degree of 'order' in the distribution of multilayered fibres within the caps does exist, however, with multilayered cell walls common in fibres adjacent to phloem elements and around the edge of the fibre cap. These findings confirm the observations, primarily in Phyllostachys viridi-glaucescens. The layering structure was not found to be specifically related to the thickness of the cell wall.     

Uneven distribution of nutrients in the root zone affects the incidence of blossom end rot and concentration of calcium and potassium in fruits of tomato

Tomato plants (Lycopersicon esculentum Mill. var. DRK) were grown hydroponically to determine the effect of an uneven distribution of nutrients in the root zone on blossom end rot (BER) and Ca and K concentrations in the fruits. The plants were grown in rockwool with their root system divided into two portions. Each portion was irrigated with nutrient solutions with either the same or the different electrical conductivity (EC) in the range 0 to 6 dS m^–1. Solutions with high EC supplied to both sides of the root system significantly increased the incidence of BER. However, when only water or a solution of low EC was supplied to one portion, BER was reduced by 80%. Fruit yields were significantly higher (P<0.01) for plants that received solutions of the uneven EC treatments (6/0 or 4.5/0 EC treatment). Plants supplied with solutions of uneven EC generally had higher leaf and fruit concentrations of Ca but lower concentrations of K than those supplied with solutions of high EC. There was no difference in Ca concentration at the distal end of young fruits of the uneven EC treatment but it was reduced in the high EC treatments. The concentration of K in the mature fruits of the uneven EC treatments was lower than that of the high EC treatments and higher or similar that of the 3/3 or 2.5/2.5 EC treatments (controls). A clear relationship was found between the incidence of BER and the exudation rate. High rate of xylem exudation was observed in the uneven EC treatments. Reduction of BER in the uneven EC treatments is most likely to be the effect of high exudation rate on Ca status in the young fruits. It was concluded that high EC of solution had positive effects on Ca concentration and incidence of BER provided that nutrient solution with low EC or water is supplied to the one portion of the root system.     

The effects of copper nutrition and developmental state on the biosynthesis of diamine oxidase in clover leaves

The biosynthesis of diamine oxidase (DAO; EC 1.4.3.6) in leaf blades of subterranean clover (Trifolium subterraneum L. cv Seaton Park) was followed by labeling whole plants with ¹⁴CO₂. A pulse-chase experiment where DAO was immunoprecipitated with anti-DAO antibodies showed that only leaf primordia and the youngest emerged leaves were able to synthesize the enzyme. The amount of DAO in young leaves of clover grown with a range of Cu treatments was determined by its enzymic activity and by single radial immunodiffusion against anti-DAO antibodies; both parameters were highly correlated with the Cu concentration of the leaf. Further, anti-DAO antibodies reacted against apo-DAO prepared in vitro indicating that apo-DAO was absent from Cu-deficient leaves. These results suggest that the biosynthesis of DAO in young clover leaves is controlled by the Cu concentrations of the leaves. Poly(A) mRNA purified from leaf primordia and young emerging leaves of plants with either a high Cu or low Cu supply was translated in wheat germ and rabbit reticulocyte cell-free systems. No differences between the two Cu treatments could be seen in the sodium dodecyl sulfate-polyacrylamide gel electrophoresis profiles of the translation products after fluorography. However, anti-DAO antibodies did not detect any DAO synthesized in vitro from either treatment.     

鼎湖山森林土壤渗透水酸度和无机氮含量对模拟氮沉降增加的早期响应

在林地分别喷加0、50、100和150kg N hm^-1a^-1,研究鼎湖山马尾松林、马尾松针阔混交林和季风常绿阔叶林中土壤20cm深渗透水酸度和无机氮含量在开始9个月的变化。结果表明,3种森林对照样方土壤渗透水pH值为3．82—4．24,外加氮处理使其平均降低了0．08—0．18。3个森林对照样方土壤渗透水无机氮平均含量分别为6．14、6．66和11．64mg L^-1,铵态氮占15．0％、11．9％和3．0％。外加氮处理使3种森林土壤渗透水铵态氮和硝态氮含量均有不同程度的提高,这表明外加氮处理不但增加了无机氮从森林土壤流失的潜力,而且使土壤进一步酸化。     

UV-C和水杨酸对虎杖叶中白藜芦醇和虎杖苷的诱导作用

以不同叶龄虎杖叶为材料,研究了不同剂量紫外线(UV-C)和10mmol/L水杨酸(SA)处理对叶中白藜芦醇和虎杖苷含量的影响。结果显示:2.4kJ/m2UV-C照射并经暗处理36h后可使虎杖嫩叶中白藜芦醇含量增加近1倍,暗处理24h后虎杖苷含量提高了30%以上,但是老叶中两者含量没有显著变化。SA处理48h后,嫩叶中白藜芦醇的含量提高约5倍,同时虎杖苷的含量下降了60%并且伴随着一个新成分的产生;老叶中白藜芦醇提高约2倍,虎杖苷的含量则没有显著变化。研究表明SA和UV-C对不同叶龄的虎杖叶片中白藜芦醇和虎杖苷合成均有诱导作用,且嫩叶比老叶表现得更为明显。     

Variation in the nutritional value of leaf and stem fractions of nineteen leucaena lines

In two experiments, 2.5-month-old edible regrowth of the genus Leucaena (varying in tolerance to the leucaena psyllid pest attack) were analysed for nutritive value. In Experiment 1, forage of nine lines was separated into young leaves, young stems (within woody fraction < 5 mm diameter), old leaves and old stems (within a woody fraction 5-10 mm diameter) whereas in Experiment 2, forage of the other 10 lines was separated in main stem leaves, branch leaves, young branches, young main stems, old main stems and old branches for analysis. Major differences were recorded between lines and forage fractions. Nutrient contents in dry matter were: nitrogen 0.7-4.8%, phosphorus 0.1-0.3%, potassium 0.2-2.5%, calcium 0.2-1.6%, magnesium 0.1-0.6%, acid detergent fibre 20-63%, lignin 8-23% and in vitro dry matter digestibility 15-69%. The leaves and immature fractions were higher in nitrogen, phosphorus, potassium and digestibility but lower in calcium, magnesium and fibre. Nutrient content in L. Leucocephala lines exceeded that in L. esculenta, L. diversifolia, leucaena hybrids and the composites. Based on the data, a management strategy that will promote young growth is suggested as appropriate for high quality forage production.     

Phloem loading strategies and water relations in trees and herbaceous plants

Most herbaceous plants employ thermodynamically active mechanisms of phloem loading, whereas in many trees, the mechanism is passive, by diffusion. Considering the different water transport characteristics of herbs and trees, we hypothesized that water relations play a role in the adoption of phloem loading strategies. We measured whole-plant hydraulic conductance (K(p)), osmolality, concentrations of polar metabolites, and key inorganic ions in recently mature leaves of 45 dicotyledonous species at midafternoon. Trees, and the few herbs that load passively, have low K(p), high osmolality, and high concentrations of transport sugars and total polar metabolites. In contrast, herbs that actively load sucrose alone have high K(p), low osmolality, and low concentrations of sugars and total polar metabolites. Solute levels are higher in sugar alcohol-transporting species, both herbs and trees, allowing them to operate at lower leaf water potentials. Polar metabolites are largely responsible for leaf osmolality above a baseline level (approximately 300 mm) contributed by ions. The results suggest that trees must offset low K(p) with high concentrations of foliar transport sugars, providing the motivating force for sugar diffusion and rendering active phloem loading unnecessary. In contrast, the high K(p) of most herbaceous plants allows them to lower sugar concentrations in leaves. This reduces inventory costs and significantly increases growth potential but necessitates active phloem loading. Viewed from this perspective, the elevation of hydraulic conductance marks a major milestone in the evolution of the herbaceous habit, not only by facilitating water transport but also by maximizing carbon use efficiency and growth.