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The arabidopsis ATHB-8 HD-zip protein acts as a differentiation-promoting transcription factor of the vascular meristems.

ATHB-8, -9, -14, -15, and IFL1/REV are members of a small homeodomain-leucine zipper family whose genes are characterized by expression in the vascular tissue. ATHB-8, a gene positively regulated by auxin (Baima et al., 1995), is considered an early marker of the procambial cells and of the cambium during vascular regeneration after wounding. Here, we demonstrate that although the formation of the vascular system is not affected in athb8 mutants, ectopic expression of ATHB-8 in Arabidopsis plants increased the production of xylem tissue. In particular, a careful anatomical analysis of the transgenic plants indicated that the overexpression of ATHB-8 promotes vascular cell differentiation. First, the procambial cells differentiated precociously into primary xylem. In addition, interfascicular cells also differentiated precociously into fibers. Finally, the transition to secondary growth, mainly producing xylem, was anticipated in transgenic inflorescence stems compared with controls. The stimulation of primary and secondary vascular cell differentiation resulted in complex modifications of the growth and development of the ATHB-8 transgenic plants. Taken together, these results are consistent with the hypothesis that ATHB-8 is a positive regulator of proliferation and differentiation, and participates in a positive feedback loop in which auxin signaling induces the expression of ATHB-8, which in turn positively modulates the activity of procambial and cambial cells to differentiate.

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Mutational analysis of branching in pea. Evidence that Rms1 and Rms5 regulate the same novel signal.

The fifth increased branching ramosus (rms) mutant, rms5, from pea (Pisum sativum), is described here for phenotype and grafting responses with four other rms mutants. Xylem sap zeatin riboside concentration and shoot auxin levels in rms5 plants have also been compared with rms1 and wild type (WT). Rms1 and Rms5 appear to act closely at the biochemical or cellular level to control branching, because branching was inhibited in reciprocal epicotyl grafts between rms5 or rms1 and WT plants, but not inhibited in reciprocal grafts between rms5 and rms1 seedlings. The weakly transgressive or slightly additive phenotype of the rms1 rms5 double mutant provides further evidence for this interaction. Like rms1, rms5 rootstocks have reduced xylem sap cytokinin concentrations, and rms5 shoots do not appear deficient in indole-3-acetic acid or 4-chloroindole-3-acetic acid. Rms1 and Rms5 are similar in their interaction with other Rms genes. Reciprocal grafting studies with rms1, rms2, and rms5, together with the fact that root xylem sap cytokinin concentrations are reduced in rms1 and rms5 and elevated in rms2 plants, indicates that Rms1 and Rms5 may control a different pathway than that controlled by Rms2. Our studies indicate that Rms1 and Rms5 may regulate a novel graft-transmissible signal involved in the control of branching.

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Nutrition of a developing legume fruit: functional economy in terms of carbon, nitrogen, water.

The economy of functioning of the developing fruit of white lupin (Lupinus albus L.) is assessed quantitatively in relation to intake and usage of carbon, nitrogen, and water. Of every 100 units of carbon imported from the parent plant, 52 are incorporated into seeds, 37 into nonmobilizable material of the pod, and the remaining 11 lost as CO(2) to the atmosphere. An illuminated fruit can make net gains of CO(2) from the atmosphere during the photoperiods of all but the last 2 weeks of its life, suggesting that it is active in assimilation of CO(2) respired from pods and seeds. This conservation activity is important to carbon economy.Phloem supplies 98% of the fruit's carbon and 89% of its nitrogen. Most of the xylem's contribution enters early in development. Xylem and phloem supply similar sets of amino compounds, amides predominating. Ninety-six per cent of the fruit's nitrogen becomes incorporated into seeds. Sixteen per cent of the seed's nitrogen is mobilized from the senescing pod.The transpiration ratio of the fruit is 22.5 ml per gram dry matter accumulated. Xylem supplies 60% of a fruit's total water requirement and the equivalent of two-thirds of its transpiration loss. Phloem becomes prominent as a water donor once the seeds start to fill.The fruit exhibits a 31% conversion by weight of organic imports into food reserves of seeds. This entails an intake through vascular channels of 1756 mg sucrose and 384 mg amino compounds and an accumulation in seeds of 412 mg protein, 132 mg oil, and 110 mg perchloric acid-soluble carbohydrate.

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Growth-induced Water Potentials in Plant Cells and Tissues.

A physical analysis of water movement through elongating soybean (Glycine max L. Merr.) hypocotyls was made to determine why significant water potentials persist in growing tissues even though the external water potentials were zero and transpiration is virtually zero. The analysis was based on a water transport theory modified for growth and assumed that water for growing cells would move through and along the cells in proportion to the conductivity of the various pathways.Water potentials calculated for individual cells were nearly in local equilibrium with the water potentials of the immediate cell surroundings during growth. However, water potentials calculated for growing tissue were 1.2 to 3.3 bars below the water potential of the vascular supply in those cells farthest from the xylem. Only cells closest to the xylem had water potentials close to that of the vascular supply. Gradients in water potential were steepest close to the xylem because all of the growth-sustaining water had to move through this part of the tissue. Average water potentials calculated for the entire growing region were -0.9 to -2.2 bars depending on the tissue diffusivity.For comparison with the calculations, average water potentials were measured in elongating soybean hypocotyls using isopiestic thermocouple psychrometers for intact and excised tissue. In plants having virtually no transpiration and growing in Vermiculite with a water potential of -0.1 bar, rapidly growing hypocotyl tissue had water potentials of -1.7 to -2.1 bars when intact and -2.5 bars when excised. In mature, nongrowing hypocotyl tissue, average water potentials were -0.4 bar regardless of whether the tissue was intact or excised.The close correspondence between predicted and measured water potentials in growing tissue indicates that significant gradients in water potential are required to move growth-associated water through and around cells over macroscopic distances. The presence of such gradients during growth indicates that cells must have different cell wall and/or osmotic properties at different positions in the tissue in order for organized growth to occur. The mathematical development used in this study represents the philosophy that would have to be followed for the application of contemporary growth theory when significant tissue water potential gradients are present.

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Allantoin and Allantoic Acid in the Nitrogen Economy of the Cowpea (Vigna unguiculata [L.] Walp.).

The ureides, allantoin and allantoic acid, represented major fractions of the soluble nitrogen pool of nodulated plants of cowpea (Vigna unguiculata [L.] Walp. cv. Caloona) throughout vegetative and reproductive growth. Stem and petioles were the principal sites of ureide accumulation, especially in early fruiting.Labeling studies using (14)CO(2) and (15)N(2) and incubation periods of 25 to 245 minutes indicated that synthesis of allantoin and allantoic acid in root nodules involved currently delivered photosynthate and recently fixed N, and that the ureides were exported from nodule to shoot via the xylem. From 60 to 80% of xylem-borne N consisted of ureides; the remainder was glutamine, asparagine, and amino acids. Allantoin predominated in the soluble N fraction of nodules and fruits, allantoin and allantoic acid were present in approximately equal proportions in xylem exudate, stems, and petioles.Extracts of the plant tissue fraction of nitrogen-fixing cowpea nodules contained glutamate synthase (EC 2.6.1.53) and glutamine synthetase (EC 6.3.1.2), but little activity of glutamate dehydrogenase (EC 1.4.1.3). High levels of uricase (EC 1.7.3.3) and allantoinase (EC 3.5.2.5) were also detected. Allantoinase but little uricase was found in extracts of leaflets, pods, and seeds.Balance sheets were constructed for production, storage, and utilization of ureide N during growth. Virtually all (average 92%) of the ureides exported from roots was metabolized on entering the shoot, the compounds being presumably used as N sources for protein synthesis.

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Economy of Carbon and Nitrogen in Nodulated and Nonnodulated (NO(3)-grown) Cowpea [Vigna unguiculata (L.) Walp.].

The response of non-nodulated cowpea (Vigna unguiculata (L.) Walp. cv Caloona) to a wide range of NO(3) levels in the rooting medium was studied 40 days after sowing by in vitro assays of plant organs for NO(3) reductase (EC 1.6.6.1) and analyses of root bleeding (xylem) sap for nitrogenous solutes. Plants fed 1, 5, 10, 20, and 40 millimolar NO(3) showed, respectively, 64, 92, 94, and 91% of their total reductase activity in shoots and 34, 30, 66, 62, and 58% of the total N of their xylem sap as NO(3). These data, and the absence in the plants of significant pools of stored NO(3), indicated that shoots were major organs of NO(3) assimilation, especially at levels of NO(3) (10 to 40 millimolar) that maintained plant growth at near maximum rates. Partitioning and utilization of C and N were studied in nodulated, minus NO(3) plants and non-nodulated plants fed 10 or 20 millimolar NO(3), the levels of NO(3) which gave rates of growth and N assimilation closest to those of the symbiotic plants. The conversion of the C of net photosynthate to dry matter was similar in nodulated plants (67%) and NO(3)-grown plants (64%), but greater proportions of photosynthate were translocated to below ground parts of nodulated plants (37%) than of NO(3)-fed plants (23 to 26%). Greater photosynthate consumption by nodulated roots was associated with proportionately greater root growth and respiration and 2-fold greater export of C in xylem than in the NO(3)-fed plants. Theoretical considerations suggest that the elevated CO(2) output of nodulated roots was due not only to CO(2) loss associated with nodule function, but also to a much greater nonassimilatory component of respiration in the supporting root of the nodulated plant compared to roots of the NO(3)-fed plants. Data are compared with previously published information from other legumes.

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Synthesis, Storage, and Utilization of Amino Compounds in White Lupin (Lupinus albus L.).

Changes in total N and in free amino compounds were followed during growth of nodulated white lupin. Leaflets contained the greatest fraction of plant N but had lower proportions (1 to 4%) of their N in soluble amino form than stem + petioles (10 to 27%) and reproductive parts (15 to 33%). Mobilization of free amino compounds from plant parts to fruits contributed at most only 7% of the total N intake of fruits, compared with 50% in mobilization of other forms of N and 43% from fixation during fruiting. Asparagine was usually the most abundant free amino compound in plant parts, followed by glutamine and alanine. Valine, glycine, isoleucine, aspartic acid and gamma-aminobutyric acid comprised the bulk of the remaining soluble amino N. Composition of tissue pools of amino-N closely resembled that of xylem and phloem exudates. Data on N flow and utilization were combined with information on composition of transport fluids to quantify syntheses, exchanges, and consumptions of asparagine, glutamine, aspartic acid, and valine by organs of the 51- to 58-day plant. These amino compounds carried 56, 29, 5, and 2%, respectively, of the N exported from nodules and contributed in roughly commensurate proportions to transport exchanges and N increments of plant parts. There were, however, more than expected involvements of glutamine and valine in mobilization of N from lower leaves, of asparagine in xylem to phloem transfer, and of aspartic acid in cycling of N through the root, and there was a less than expected participation of aspartic acid in xylem to phloem transfer and in phloem translocation to the shoot apex. The significance of these differences is discussed.

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Potassium nutrition and translocation in sugar beet.

The effect of increased net foliar K(+) accumulation on translocation of carbon was studied in sugar beet (Beta vulgaris, L. var. Klein E and US H20) plants. Net accumulation of recently absorbed K(+) was studied by observing arrival of (42)K(+) per unit area of leaf. Labeled K(+) was added to give an initial concentration at 2 or 10 millimolar K(+) in mineral nutrient solution. Because the newly arrived K(+) constitutes a small part of the total leaf K(+) in plants raised in 10 millimolar K(+), export of (42)K(+) by phloem was negligible over the 2- to 3-day period; consequently, accumulation is a measure of arrival in the xylem. In leaves from plants in 2 millimolar K(+), export by the phloem was estimated to be of the same order as import by the xylem; K(+) per area was observed to remain at a steady-state level. Increasing the supply of K(+) to 10 millimolar caused arrival in the xylem to increase 2- to 3-fold; K(+) per area increased gradually in the mature leaves. Neither net carbon exchange nor translocation of sugar increased in response to a faster rate of arrival of K(+) over a 6- to 8-hour period. In the absence of short-term effects, it is suggested that K(+)-promoted increase in synthetic metabolism may be the basis of the increased carbon assimilation and translocation in plants supplied with an above-minimal level of K(+).

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An interpretation of some whole plant water transport phenomena.

A treatment of water flow into and through plants to the evaporating surface of the leaves is presented. The model is driven by evaporation from the cell wall matrix of the leaves. The adsorptive and pressure components of the cell wall matric potential are analyzed and the continuity between the pressure component and the liquid tension in the xylem established. Continuity of these potential components allows linking of a root transport function, driven by the tension in the xylem, to the leaf water potential. The root component of the overall model allows for the solvent-solute interactions characteristic of a membrane-bound system and discussion of the interactions of environmental variables such as root temperature and soil water potentials. A partition function is developed from data in the literature which describes how water absorbed by the plant might be divided between transpiration and leaf growth over a range of leaf water potentials.Relationships between the overall system conductance and the conductance coefficients of the various plant parts (roots, xylem, leaf matrix) are established and the influence of each of these discussed.The whole plant flow model coupled to the partition function is used to simulate several possible relationships between leaf water potential and transpiration rate. The effects of changing some of the partition function coefficients, as well as the root medium water potential on these simulations is illustrated.In addition to the general usefulness of the model and its ability to describe a wide range of situations, we conclude that the relationships used, dealing with bulk fluid flow, diffusion, and solute transport, are adequate to describe the system and that analogically based theoretical systems, such as the Ohm's law analogy, probably ought to be abandoned for this purpose.

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Spontaneous Phloem bleeding from cryopunctured fruits of a ureide-producing legume.

The vasculature of the dorsal suture of cowpea (Vigna unguiculata [L.] Walp) fruits bled a sugar-rich exudate when punctured with a fine needle previously cooled in liquid N(2). Bleeding continued for many days at rates equivalent to 10% of the estimated current sugar intake of the fruit. A phloem origin for the exudate was suggested from its high levels (0.4-0.8 millimoles per milliliter) of sugar (98% of this as sucrose) and its high K(+) content and high ratio of Mg(2+) to Ca(2+). Fruit cryopuncture sap became labeled with (14)C following feeding of [(14)C]urea to leaves or adjacent walls of the fruit, of (14)CO(2) to the pod gas space, and of [(14)C] asparagine or [(14)C]allantoin to leaflets or cut shoots through the xylem. Rates of translocation of (14)C-assimilates from a fed leaf to the puncture site on a subtended fruit were 21 to 38 centimeters per hour. Analysis of (14)C distribution in phloem sap suggested that [(14)C]allantoin was metabolized to a greater extent in its passage to the fruit than was [(14)C] asparagine. Amino acid:ureide:nitrate ratios (nitrogen weight basis) of NO(3)-fed, non-nodulated plants were 20:2:78 in root bleeding xylem sap versus 90:10:0.1 for fruit phloem sap, suggesting that the shoot utilized NO(3)-nitrogen to synthesize amino acids prior to phloem transfer of nitrogen to the fruit. Feeding of (15)NO(3) to roots substantiated this conclusion. The amino acid:ureide ratio (nitrogen weight basis) of root xylem sap of symbiotic plants was 23:77 versus 89:11 for corresponding fruit phloem sap indicating intense metabolic transfer of ureide-nitrogen to amino acids by vegetative parts of the plant.

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Effects of n(2) deficiency on transport and partitioning of C and N in a nodulated legume.

Nodulated root systems of white lupin (Lupinus albus L. cv Ultra: Rhizobium strain WU425) were exposed to Ar:O(2) (80:20, v/v) or Ar:N(2):O(2) (70:10:20, v/v/v) and C and N partitioning were examined over a 9- or 10-day period in comparison with control plants with nodulated roots retained in air. Accumulation of N ceased in plants exposed to Ar:O(2) or was much reduced in plants exposed to Ar:N(2):O(2), but net C assimilation rates and profiles of C utilization remained similar to those of control N(2)-fixing plants. There was, however, a proportional reduction in CO(2) evolution from nodulated roots of the Ar:O(2) treatment. Xylem N levels fell rapidly after application of Ar:O(2). C:N ratios of phloem sap of petioles and of stem base rose during the first day of Ar:O(2) treatment and then fell progressively back to levels close to that of control plants as leaf reserves of N became available for loading of phloem. Stem top phloem sap increased progressively in C:N ratio throughout Ar:O(2) treatment, presumably due to increasing shortage of xylem derived N for xylem to phloem exchange. Reexposure of Ar:O(2)-treated nodulated root systems to air prompted a rapid recovery of N(2) fixation and restoration of plant N status. Rates of N(2) fixation in plants whose roots were exposed to a range of N(2) concentrations indicated an apparent K(m) of 10% N(2) for the attached intact white lupin nodule.

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Economy of water, carbon, and nitrogen in the developing cowpea fruit.

The nutritional economy of the fruit of cowpea (Vigna unguiculata (L.) Walp cv Vita 3) was assessed quantitatively from intake and utilization of carbon, nitrogen, and water. Fruits failed to make net gains of CO(2) from the atmosphere during daytime, although pod photosynthesis did play a role in the fruit's carbon economy by refixing a proportion of the fruit's respired CO(2). Of every 100 units by weight of carbon entering the fruit, 70.4 were finally incorporated into seeds, 10.3 remained as nonmobilizable material in pod walls, and the remaining 19.3 were lost in fruit respiration. Phloem supplied 97% of the fruit's carbon and 72% of its nitrogen. The xylem contribution of nitrogen occurred mainly in early growth. Ninety-six% of the fruit's nitrogen was incorporated into seeds, approximately 10% of this mobilized from the senescing pod. The mean transpiration ratio of the fruit was very low-8 milliliters water transpired per gram dry matter accumulated. Models of carbon, nitrogen, and water flow were constructed for the two consecutive 11 day periods of fruit development, and indicated a considerably greater entry of water through xylem and phloem than could be accounted for in changes in fruit tissue water and transpiration loss. This discrepancy was greater in the second half of fruit growth and was interpreted as evidence that a significant fraction of the water entering the fruit through phloem cycled back to the parent plant via the xylem.

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Quantifying Apoplastic Flux through Red Pine Root Systems Using Trisodium, 3-hydroxy-5,8,10-pyrenetrisulfonate.

The fluorescent compound trisodium, 3-hydroxy-5,8,10-pyrenetrisulfonate (PTS) was used to quantify the apoplastic flux through red pine (Pinus resinosa Ait.) root systems-that portion of the total water flux reaching the xylem without ever crossing a semipermeable membrane. Flow was induced by pressure through detopped root systems, and by transpiration through intact seedlings. Apoplastic flux was determined by multiplying total flux by the ratio of PTS concentration in the xylem exudate to PTS concentration in the bathing medium.Under aeration, apoplastic flux was less than 1% of total flux. Under anaerobic conditions, up to 50% of total flux was apoplastic suggesting that anaerobic conditions change the pathway of water flow into root xylem. The change under anaerobic conditions was reversible. Detopped root systems under pressure and intact seedlings under transpiration gave similar results. In detopped root systems, the magnitude of the pressure gradient may alter the apoplastic contribution to total flux.

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p-Fluorophenylalanine-Induced Restriction of Ion Uptake and Assimilation by Maize Roots.

Roots of decapitated maize seedlings (Zea mays L.) were exposed for 12 hours to 1.0 millimolar KNO(3) (98.5 atom per cent (15)N) in the presence and absence (control) of 0.1 millimolar p-fluorophenylalanine (FPA), an analog of the amino acid phenylalanine. FPA decreased nitrate uptake but had little effect on potassium uptake. In contrast, accumulation of both ions in the xylem exudate was greatly restricted. The proportion of reduced (15)N-nitrogen that was translocated at each time was also restricted by FPA. These observations are interpreted as indicating that synthesis of functional protein(s) is required for nitrate uptake and for transport of potassium, nitrate, and reduced-(15)N from xylem parenchyma cells into xylem elements. The effect of FPA on nitrate reduction is less clear. Initially, FPA limited nitrate reduction more than nitrate uptake, but by 8 hours the cumulative reduction of entering nitrate was similar ( approximately 35%) in both control and FPA-treated roots. A relationship between nitrate uptake and nitrate reduction is implied. It is suggested that nitrate influx regulates the proportion of nitrate reductase in the active state, and thereby regulates concurrent nitrate reduction in decapitated maize seedlings.

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Measurement of the water potential of stored potato tubers.

A method of measuring the water potential of stored potato tubers (Solanum tuberosum L.) was needed to investigate the relationship of bacterial soft rot in tubers to water potential. Pressure chamber measurements, while useful for tubers with functional stolons, cannot be made on stored tubers. Measurements could be made on excised tissue pieces in a hygrometer chamber and with hygrometers implanted into tubers. We report here our evaluation of these hygrometric methods using a comparison with the pressure chamber on tubers harvested with stolons intact.In tubers of high water potential, measurements on excised tissue were as much as 0.5 megapascals lower than the pressure chamber, probably due to turgor-driven expansion of the sample when freed from constraints imposed by surrounding tissue. Good agreement (+/-0.05 megapascals) was found between the implanted hygrometer and the pressure chamber at potentials higher than -0.5 megapascals. At lower water potentials, both hygrometer measurements were higher than the pressure chamber. Respirational heating of the tissue contributed to the increase in the excised tissue samples, but not with the implanted hygrometers because of the hygrometer design. The osmotic pressure balanced the pressure chamber measurement of potential at -0.7 megapascals, but was too small to do so at lower potentials. At most, 25% of this discrepancy can be accounted for by dilution by apoplastic water. We believe that the pressure chamber measurement is too low at low water potentials and that the error is associated with air bubbles in the xylem. At low potentials air emerged from xylem vessels along with sap, and fewer xylem emitted sap as potentials decreased.

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Mechanism of export of organic material from the developing fruits of pea.

Pisum sativum L. fruits export a small quantity of radiolabeled substances to other plant parts after the fruits are allowed to photosynthesize in the presence of (14)CO(2). Export was uninhibited by peduncle girdling suggesting an apoplastic route for transport of material, presumably by ;reverse' flow in the peduncle xylem. To determine if any diurnal water potential gradient formed between pea leaves and fruit might be responsible for the observed export, the water potentials of the various organs were monitored over 24 hours. Water potential differences of up to 7.5 bars existed between leaves and fruit in long photoperiods, and up to 2.5 bars in short photoperiods. Pulses of (14)CO(2) labeling indicated that initial delivery of exported label was to ;transpirational sinks,' with subsequent redistribution of label to metabolic sinks. Export to the apical bud appeared to be direct via the xylem. Application of membrane-impermeable inulin to a surgically opened seed coat ;cup' resulted in export mainly to the subtending leaf with little redistribution. Simultaneous application of sucrose to the seed coat resulted in more extensive distribution of the sucrose, consistent with reloading of the sucrose into mature leaf phloem. Thus, export of material from fruits appears to occur via a xylem pathway in response to transpirationally derived water potential gradients.

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Inhibition of nodule functioning in cowpea by a xanthine oxidoreductase inhibitor, allopurinol.

Allopurinol (1H-pyrazolo-[3,4-d]pyrimidine-4-ol), an inhibitor of xanthine oxidation in ureide-producing nodulated legumes, was taken up from the rooting medium, translocated in xylem, and transferred to nodules of both the ureide-forming cowpea (Vigna unguiculata L. Walp.) and the amide-forming white lupin (Lupinus albus L.). Cowpea suffered severe nitrogen deficiency, extreme chlorosis, and reduced growth, whereas lupin was unaffected by the inhibitor. Similar results were obtained with oxypurinol (1H-pyrazolo-[3,4-d]pyrimidine-4,6-diol). Xylem composition of symbiotic cowpea was markedly changed by allopurinol. Ureides fell to a very low level, but xanthine and, to a lesser extent, hypoxanthine increased markedly. Xylem glutamine was also reduced, but there was little change in other amino acids. Nitrogenase (EC 1.7.99.2) activity of intact nodulated plants or nodulated root segments of plants treated with allopurinol or oxypurinol for 24 hours or more was severely inhibited in cowpea but unaffected in lupin for periods of exposure up to 9 days. Nitrogenase activity of slices of nodules prepared from allopurinol-treated cowpea showed inhibition comparable to that of intact plants. Breis prepared from nodules of treated plants showed no reduction in nitrogenase, nor was there reduction in activity of breis following addition of allopurinol, xanthine, or a range of purine pathway intermediates. Increasing the O(2) concentration in assays above 20% (volume/volume) reversed inhibition of nitrogenase by allopurinol in intact nodulated roots. It was concluded for cowpea that allopurinol not only inhibited ureide synthesis but also caused inhibition of nitrogenase activity, thereby leading to progressive dysfunction and eventual senescence of nodules. The mechanistic relationships between inhibition of ureide biosynthesis, changes in gaseous diffusion resistance, and reduced nitrogenase activity remain obscure.

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Water Transport in the Liana Bauhinia fassoglensis (Fabaceae).

To determine the efficiency of xylem conductance in the liana (woody vine) Bauhinia fassoglensis Kotschy ex Schweinf., we measured hydraulic conductance per unit stem length (measured K(h)), leaf-specific conductivity (LSC = K(h)/distal leaf area), transpiration rate (E), xylem water potential (epsilon), vessel number, and vessel diameter. The measured K(h) was 49% (se = 7%) of the predicted K(h) from Poiseuille's law. The mean LSC for unbranched stem segments was 1.10 x 10(-8) square meters per megapascal per second (se = 0.07). LSCs were much lower (about 0.2) at branch junctions. At midday, with E at 7 x 10(-8) meters per second, the measured drop in epsilon was about 0.08 megapascal per meter along the stems and branches and about 0.27 megapascal in going from stem to leaf. In addition, there was a drop of about 0.20 megapascal at branch junctions as predicted by E/LSC. In diurnal measurements leaf epsilon never dropped below about -1.2 megapascal. For long (e.g. 16 meters) stems, the predicted mid-day drop in epsilon through the xylem transport system might be great enough to have substantial physiological impact.

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