Biomedical subjects

D B Layzell

Publications and source records attributed to D B Layzell.

30 records · Page 2Linked to original sources

PubMed1985

Effect of N Source on the Steady State Growth and N Assimilation of P-limited Anabaena flos-aquae.

Phosphate-limited chemostat cultures were used to study cell growth and N assimilation in Anabaena flos-aquae under various N sources to determine the relative energetic costs associated with the assimilation of NH(3), NO(3) (-), or N(2). Expressed as a function of relative growth rate, steady state cellular P contents and PO(4) assimilation rates did not vary with N-source. However, N-source did alter the maximal PO(4)-limited growth rate achieved by the cultures: the NO(3) (-) and N(2) cultures attained only 97 and 80%, respectively, of the maximal growth rate of the NH(3) grown cells. Cellular biomass and C contents did not vary with growth rate, but changed with N source. The NO(3) (-)-grown cells were the smallest (627 +/- 34 micromoles C . 10(-9) cells), while NH(3)-grown cells were largest (900 +/- 44 micromoles C . 10(-9) cells) and N(2)-fixing cells were intermediate (726 +/- 48 micromoles C . 10(-9) cells) in size. In the NO(3) (-)-and N(2)-grown cultures, N content per cell was only 57 and 63%, respectively, of that in the NH(3)-grown cells. Heterocysts were absent in NH(3)-grown cultures but were present in both the N(2) and NO(3) (-) cultures. In the NO(3) (-)-grown cultures C(2)H(2) reduction was detected only at high growth rates, where it was estimated to account for a maximum of 6% of the N assimilated. In the N(2)-fixing cultures the acetylene:N(2) ratio varied from 3.4:1 at lower growth rates to 3.0:1 at growth rates approaching maximal.Compared with NH(3), the assimilation of NO(3) (-) and N(2) resulted either in a decrease in cellular C (NO(3) (-) and N(2) cultures) or in a lower maximal growth rate (N(2) culture only). The observed changes in cell C content were used to calculate the net cost (in electron pair equivalents) associated with growth on NO(3) (-) or N(2) compared with NH(3).

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PubMed1985

Modeling the C Economy of Anabaena flos-aquae: Estimates of Establishment, Maintenance, and Active Costs Associated with Growth on NH(3), NO(3), and N(2).

Steady state cultures of Anabaena flos-aquae were established over a wide range of phosphate-limited growth rates while N was supplied as either NH(3), NO(3) (-), or N(2) gas. At growth rates greater than 0.03 per hour, rates of gross and net carbon fixation were similar on all N sources. However, at lower growth rates (<0.03 per hour) in the NO(3) (-) and N(2) cultures, gross photosynthesis greatly exceeded net photosynthesis. The increase in photosynthetic O(2) evolution with growth rate was greatest when N requirements were met by NO(3) (-) and least when met by NH(3). These results were combined with previously reported measurements of cellular chemical composition, N assimilation, and acetylene reduction (Layzell, Turpin, Elrifi 1985 Plant Physiol 78: 739-745) to construct empirical models of carbon and energy flow for cultures grown at 30, 60, and 100% of their maximal growth rate on all N sources. The models suggested that over this growth range, 89 to 100% of photodriven electrons were allocated to biomass production in the NH(3) cells, whereas only 49 to 74% and 54 to 90% were partitioned to biomass in the NO(3) (-)-and N(2)-grown cells, respectively. The models were used to estimate the relative contribution of active, maintenance, and establishment costs associated with NO(3) (-) and N(2) assimilation over the entire range of growth rates. The models showed that the relative contribution of the component costs of N assimilation were growth rate dependent. At higher growth rates, the major costs for NO(3) (-) assimilation were the active costs, while in N(2)-fixing cultures the major energetic requirements were those associated with heterocyst establishment and maintenance. It was concluded that compared with NO(3) (-) assimilation, N(2) fixation was energetically unfavorable due to the costs of heterocyst establishment and maintenance, rather than the active costs of N(2) assimilation.

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PubMed1984

A highly sensitive, flow through h(2) gas analyzer for use in nitrogen fixation studies.

Studies of H(2) evolution by N(2) fixing systems are frequently limited by an inability to accurately measure H(2) gas concentrations of less than about 10 microliters per liter. In this study, a H(2) gas analyzer is described which is able to accurately and reproducibly detect up to 100 times lower H(2) concentrations than most thermal conductivity gas chromatographs or other conventional instruments used for the measurement of H(2) gas. This high level of sensitivity (maximum of about 0.02 microliter per liter H(2) per millivolt output) and the ability to continuously monitor H(2) concentration directly in a flowing gas stream, makes this instrument well suited for use in an open gas exchange system.Since the sensor used in the instrument was also sensitive to other combustible gases, it was necessary to demonstrate that H(2) was the only combustible gas produced by the N(2) fixing system being studied. When an air stream was passed through a pot containing nodulated soybean (Glycine max L.) roots, gas chromatographic analysis of the effluent gas stream revealed that H(2) was the only combustible gas present. These results were supported by other studies in which no combustible gases were detected in the effluent gas stream from soybean roots nodulated with USDA 110, a Rhizobium strain which displays active uptake hydrogenase activity.

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PubMed1984

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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PubMed1982

Modeling C and N transport to developing soybean fruits.

Xylem sap and phloem exudates from detached leaves and fruit tips were collected and analyzed during early pod-fill in nodulated soybeans (Glycine max (L.) Merr. cv Wilkin) grown without (-N) and with (+N) NH(4)NO(3). Ureides were the predominant from (91%) of N transported in the xylem of -N plants, while amides (45%) and nitrate (23%) accounted for most of the N in the xylem of +N plants. Amino acids (44%) and ureides (36%) were the major N forms exported in phloem from leaves in -N plants, but amides (63%) were most important in +N plants. Based on the composition of fruit tip phloem, ureides (55% and 33%) and amides (26% and 47%) accounted for the majority of N imported by fruits of -N and +N plants, respectively.C:N weight ratios were lowest in xylem exudate (1.37 and 1.32), highest in petiole phloem (24.5 and 26.0), and intermediate in fruit tip exudate (12.6 and 12.1) for the -N and +N treatments, respectively. The ratios were combined with data on fruit growth and respiration to construct a model of C and N transport to developing fruits. The model indicates xylem to phloem transfer provides 35% to 52% of fruit N. Results suggest the phloem entering fruits oversupplies their N requirement so that 13% of the N imported is exported from fruit in the xylem.

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PubMed1981

Partitioning of carbon and nitrogen and the nutrition of root and shoot apex in a nodulated legume.

Empirically based models depicting exchanges of C, N, and H(2)O in phloem and xylem among organs of nodulated white lupin (Lupinus albus cv Ultra) were constructed for the interval 51 to 58 days after sowing. Information was incorporated on the economy of C, N, and H(2)O in plant parts, the solute composition of transport fluids collected at selected sites on the plant, and the photosynthetic inputs, transpirational losses, and translocatory activities of different age groups of leaflets and stem + petiole segments of the shoot. Partitioning of C and N showed preferential transfer of N to the shoot apex, which imported 13 milligrams C per milligram N, compared with 54 milligrams C per milligram N for the nodulated root. Leaves translocated assimilates at a C:N weight ratio of 43 to 59, and older leaves serving the roots produced the translocate most rich in N relative to C. The shoot apex was enriched with N, additional to its intake from leaves, by direct uptake of xylem fluid (C:N ratio, 2.4) and receipt of nitrogenous solutes transferred from xylem to upward-moving phloem streams in upper regions of the stem. The models for flow of N and H(2)O indicated that xylem streams passing to leaves were substantially less rich in N than the adjacent stream moving through the body of the stem and that a progressive increase in concentration of N occurred within stem xylem elements from base to top of the shoot. This apparently resulted from an abstraction of N from xylem of departing leaf traces, possibly by xylem transfer cells, and a subsequent feedback of this N to xylem streams passing on up the shoot. Upper leaves and shoot apex, therefore, acquired more N from xylem per unit of H(2)O transpired than lower parts of the shoot.

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PubMed1981

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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PubMed1979

Modeling the transport and utilization of carbon and nitrogen in a nodulated legume.

An empirical modeling technique was developed for depicting quantitatively the transport and partitioning of photosynthetically fixed C and symbiotically fixed N during 10-day intervals of a 40-day period in the growth of nodulated plants of white lupin (Lupinus albus L. cv. Ultra). Model construction utilized data for C and N consumption of plant parts and C:N weight ratios of the xylem and phloem fluids serving specific plant organs. Formulas were derived from calculating the net transport of C and N between plant parts in xylem and phloem. The models provided quantitative information on the dependence of growing organs on xylem and phloem for their supply of C and N, the cycling of N through leaflets and of C through nodules, the extent of direct incorporation of fixed N into growing nodules, and the involvement of N from shoot translocate in the nutrition of the nodulated root. Stem plus petioles abstracted considerably more N from xylem than expected from their transpirational activity. Xylem to phloem transfer of recently fixed N in mature stem and petioles was substantiated by the models, being depicted as a device for dispensing N to growing parts of the shoot extra to that attracted transpirationally in xylem or received as translocate from leaflets.

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PubMed1979

Transport of organic solutes in Phloem and xylem of a nodulated legume.

Collections of xylem exudate of root stumps or detached nodules, and of phloem bleeding sap from stems, petioles, and fruits were made from variously aged plants of Lupinus albus L. relying on nodules for their N supply. Sucrose was the major organic solute of phloem, asparagine, glutamine, serine, aspartic acid, valine, lysine, isoleucine, and leucine, the principal N solutes of both xylem and phloem. Xylem sap exhibited higher relative proportions of asparagine, glutamine and aspartic acid than phloem sap, but lower proportions of other amino acids. Phloem sap of petioles was less concentrated in asparagine and glutamine but richer in sucrose than was phloem sap of stem and fruit, suggesting that sucrose was unloaded from phloem and amides added to phloem as translocate passed through stems to sinks of the plant. Evidence was obtained of loading of histidine, lysine, threonine, serine, leucine and valine onto phloem of stems but the amounts involved were small compared with amides. Analyses of petiole phloem sap from different age groups of leaves indicated ontogenetic changes and effects of position on a shoot on relative rates of export of sucrose and N solutes. Diurnal fluctuations were demonstrated in relative rates of loading of sucrose and N solutes onto phloem of leaves. Daily variations in the ability of stem tissue to load N onto phloem streams were of lesser amplitude than, or out of phase with fluctuations in translocation of N from leaves. Data were related to recent information on C and N transport in the species.

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PubMed1979

Economy of Photosynthate Use in Nitrogen-fixing Legume Nodules: Observations on Two Contrasting Symbioses.

The economy of C use by root nodules was examined in two symbioses, Vigna unguiculata (L.) Walp. (cv. Caloona):Rhizobium CB756 and Lupinus albus L. (cv. Ultra):Rhizobium WU425 over a 2-week period in early vegetative growth. Plants were grown in minus N water culture with cuvettes attached to the nodulated zone of their primary roots for collection of evolved CO(2) and H(2). Increments in total plant N and in C and N of nodules, and C:N weight ratios of xylem and phloem exudates were studied by periodic sampling from the plant populations. Itemized budgets were constructed for the partitioning and utilization of C in the two species. For each milligram N fixed and assimilated by the cowpea association, 1.54 +/- 0.26 (standard error) milligrams C as CO(2) and negligible H(2) were evolved and 3.11 milligrams of translocated C utilized by the nodules. Comparable values for nodules of the lupin association were 3.64 +/- 0.28 milligrams C as CO(2), 0.22 +/- 0.05 milligrams H(2), and 6.58 milligrams C. More efficient use of C by cowpea nodules was due to a lesser requirement of C for synthesis of exported N compounds, a smaller allocation of C to nodule dry matter, and a lower evolution of CO(2). The activity of phosphoenolpyruvate carboxylase in nodule extracts and the rate of (14)CO(2) fixation by detached nodules were greater for the cowpea symbiosis (0.56 +/- 0.06 and 0.22 milligrams C as CO(2) fixed per gram fresh weight per hour, respectively) than for the lupin 0.06 +/- 0.02 and 0.01 milligrams C as CO(2) fixed per gram fresh weight per hour. The significance of the data was discussed in relation to current information on theoretical costs of nitrogenase functioning and associated nodule processes.

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PubMed1979

Assimilation and Transport of Nitrogen in Nonnodulated (NO(3)-grown) Lupinus albus L.

The response of nonnodulated white lupin (Lupinus albus L. cv. Ultra) plants to a range of NO(3) levels in the rooting medium was studied by in vitro assays of extracts of plant parts for NO(3) reductase (EC 1.6.6.1) activity, measurements of NO(3)-N in plant organs, and solute analyses of root bleeding (xylem) sap and phloem sap from stems and petioles. Plants were grown for 65 days with 5 millimolar NO(3) followed by 10 days with 1, 5, 15, or 30 millimolar NO(3). NO(3) reductase was substrate-induced in all tissues. Roots contained 76, 68, 62 and 31% of the total NO(3) reductase activity of plants fed with 1, 5, 15, and 30 millimolar NO(3), respectively. Stem, petioles, and leaflets contained virtually all of the NO(3) reductase activity of a shoot, the activity in extracts of fruits amounting to less than 0.3% of the total enzyme recovered from the plant. Xylem sap from NO(3)-grown nonnodulated plants contained the same organic solutes as from nodulated plants grown in the absence of combined N. Asparagine accounted for 50 to 70% and glutamine 10 to 20% of the xylem-borne N. The level of NO(3) in xylem sap amounted to 4, 13, 12, and 17% of the total xylem N at 1, 5, 15, and 30 millimolar NO(3), respectively. Xylem to phloem transfer of N appeared to be quantitatively important in supplying fruits and vegetative apices with reduced N, especially at low levels of applied NO(3). NO(3) failed to transfer in any quantity from xylem to phloem, representing less than 0.3% of the phloem-borne N at all levels of applied NO(3). Shoot organs were ineffective in storing NO(3). Even when NO(3) was supplied in great excess (30 millimolar level) it accounted for only 8% of the total N of stem and petioles, and only 2 and 1% of the N of leaflets and fruits, respectively.

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PubMed1979

Economy of Carbon and Nitrogen in a Nodulated and Nonnodulated (NO(3)-grown) Legume.

Partitioning and utilization of assimilated C and N were compared in nonnodulated, NO(3)-fed and nodulated, N(2)-fed plants of white lupin (Lupinus albus L.). The NO(3) regime used (5 millimolar NO(3)) promoted closely similar rates of growth and N assimilation as in the symbiotic plants. Over 90% of the N absorbed by the NO(3)-fed plants was judged to be reduced in roots. Empirically based models of C and N flow demonstrated that patterns of incorporation of C and N into dry matter and exchange of C and N among plant parts were essentially similar in the two forms of nutrition. NO(3)-fed and N(2)-fed plants transported similar types and proportions of organic solutes in xylem and phloem. Withdrawal of NO(3) supply from NO(3)-fed plants led to substantial changes in assimilate partitioning, particularly in increased translocation of N from shoot to root. Nodulated plants showed a lower (57%) conversion of C or net photosynthate to dry matter than did NO(3)-fed plants (69%), and their stems were only half as effective as those of NO(3)-fed plants in xylem to phloem transfer of N supplied from the root. Below-ground parts of symbiotic plants consumed a larger share (58%) of the plants' net photosynthate than did NO(3)-fed roots (50%), thus reflecting a higher CO(2) loss per unit of N assimilated (10.2 milligrams C/milligram N) by the nodulated root than by the root of the NO(3)-fed plant (8.1 milligrams C/milligram N). Theoretical considerations indicated that the greater CO(2) output of the nodulated root involved a slightly greater expenditure for N(2) than for NO(3) assimilation, a small extra cost due to growth and maintenance of nodule tissue, and a considerably greater nonassimilatory component of respiration in root tissue of the symbiotic plant than in the root of the NO(3)-fed plant.

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