Influence of diet on the distribution of nitrogen isotopes in animals

The influence of diet on the distribution of nitrogen isotopes in animals was investigated by analyzing animals grown in the laboratory on diets of constant nitrogen isotopic composition. 
The isotopic composition of the nitrogen in an animal reflects the nitrogen isotopic composition of its diet. The δ^(15)N values of the whole bodies of animals are usually more positive than those of their diets. Different individuals of a species raised on the same diet can have significantly different δ^(15)N values. The variability of the relationship between the δ^(15)N values of animals and their diets is greater for different species raised on the same diet than for the same species raised on different diets. Different tissues of mice are also enriched in ^(15)N relative to the diet, with the difference between the δ^(15)N values of a tissue and the diet depending on both the kind of tissue and the diet involved. The δ^(15)N values of collagen and chitin, biochemical components that are often preserved in fossil animal remains, are also related to the δ^(15)N value of the diet. 
The dependence of the δ^(15)N values of whole animals and their tissues and biochemical components on the δ^(15)N value of diet indicates that the isotopic composition of animal nitrogen can be used to obtain information about an animal's diet if its potential food sources had different δ^(15)N values. The nitrogen isotopic method of dietary analysis probably can be used to estimate the relative use of legumes vs non-legumes or of aquatic vs terrestrial organisms as food sources for extant and fossil animals. However, the method probably will not be applicable in those modern ecosystems in which the use of chemical fertilizers has influenced the distribution of nitrogen isotopes in food sources. 
The isotopic method of dietary analysis was used to reconstruct changes in the diet of the human population that occupied the Tehuacan Valley of Mexico over a 7000 yr span. Variations in the δ^(15)C and δ^(15)N values of bone collagen suggest that C_4 and/or CAM plants (presumably mostly corn) and legumes (presumably mostly beans) were introduced into the diet much earlier than suggested by conventional archaeological analysis.

Influence of Diet On the Distribtion of Nitrogen Isotopes in Animals

Virtually all nonequilibrium electron transfers on Earth are driven by a set of nanobiological machines composed largely of multimeric protein complexes associated with a small number of prosthetic groups. These machines evolved exclusively in microbes early in our planet's history yet, despite their antiquity, are highly conserved. Hence, although there is enormous genetic diversity in nature, there remains a relatively stable set of core genes coding for the major redox reactions essential for life and biogeochemical cycles. These genes created and coevolved with biogeochemical cycles and were passed from microbe to microbe primarily by horizontal gene transfer. A major challenge in the coming decades is to understand how these machines evolved, how they work, and the processes that control their activity on both molecular and planetary scales.

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The Microbial Engines That Drive Earth's Biogeochemical Cycles

▪ Abstract The use of stable isotope techniques in plant ecological research has grown steadily during the past two decades. This trend will continue as investigators realize that stable isotopes can serve as valuable nonradioactive tracers and nondestructive integrators of how plants today and in the past have interacted with and responded to their abiotic and biotic environments. At the center of nearly all plant ecological research which has made use of stable isotope methods are the notions of interactions and the resources that mediate or influence them. Our review, therefore, highlights recent advances in plant ecology that have embraced these notions, particularly at different spatial and temporal scales. Specifically, we review how isotope measurements associated with the critical plant resources carbon, water, and nitrogen have helped deepen our understanding of plant-resource acquisition, plant interactions with other organisms, and the role of plants in ecosystem studies. Where possible we also...

Stable Isotopes in Plant Ecology

Biological N2 fixation represents the major source of N input in agricultural soils including those in arid regions. The major N2-fixing systems are the symbiotic systems, which can play a significant role in improving the fertility and productivity of low-N soils. The Rhizobium-legume symbioses have received most attention and have been examined extensively. The behavior of some N2-fixing systems under severe environmental conditions such as salt stress, drought stress, acidity, alkalinity, nutrient deficiency, fertilizers, heavy metals, and pesticides is reviewed. These major stress factors suppress the growth and symbiotic characteristics of most rhizobia; however, several strains, distributed among various species of rhizobia, are tolerant to stress effects. Some strains of rhizobia form effective (N2-fixing) symbioses with their host legumes under salt, heat, and acid stresses, and can sometimes do so under the effect of heavy metals. Reclamation and improvement of the fertility of arid lands by application of organic (manure and sewage sludge) and inorganic (synthetic) fertilizers are expensive and can be a source of pollution. The Rhizobium-legume (herb or tree) symbiosis is suggested to be the ideal solution to the improvement of soil fertility and the rehabilitation of arid lands and is an important direction for future research.

Rhizobium-Legume Symbiosis and Nitrogen Fixation under Severe Conditions and in an Arid Climate

Reproductive as well as vegetative parameters of mature soybean (Glycine max [L.] Merr. cv. Wye) plants grown in chambers in which the aerial portion was exposed to altered pO(2) during all or part of the growth cycle were measured. Oxygen concentration was found to be a key factor controlling all phases of reproductive development. Exposure to 5% O(2) from early seedling stage to senescence increased leaf, stem, and root dry weights and reduced seed yields when compared to 21% O(2); exposure to low O(2) during the vegetative growth stage from early seedling to mid-flowering arrested pod but not seed development; exposure from mid-flowering to mid-pod filling almost completely arrested seed but not pod development; exposure from mid-pod filling to senescence arrested seed development at the mid-filling stage.Exposures to 5% O(2) initiated at mid-flowering for 1, 2, 3, 5, 10, and 15 days had no effect on seed development when the exposure was up to 3 days and produced almost total arrest when the exposure was 10 or more days, suggesting reversibility. The requirement for O(2) in seed development is independent of CO(2) concentration with similar results produced by subambient O(2) combined with ambient CO(2), elevated CO(2) up to 2000 mul/l or depressed levels of CO(2) with the CO(2)/O(2) ratio as in air. An elevated O(2) atmosphere containing 40% O(2) and ambient or elevated CO(2) inhibited total growth but did not affect the balance of vegetative to reproductive growth.We conclude that an unknown reaction or process requiring at least atmospheric concentrations of O(2) but independent of CO(2) in contrast to photorespiration is necessary for optimization of all phases of reproductive growth and the effect is reversible for exposures of up to 3 days but not for exposures of 10 days or more. We propose that this O(2) phenomenon may be the result of a unique physical process or chemical reaction associated with translocation and accumulation of assimilates in reproductive structures.

Reproductive Growth and Dry Matter Production of Glycine max (L.) Merr. in Response to Oxygen Concentration.

The N2-fixing legume nodule requires O2 for ATP production; however, the O2 sensitivity of nitrogenase dictates a requirement for a low pO2 inside the nodule. The effects of long term exposures to various pO2s on N2[C2H2] fixation were evaluated with intact soybean (Glycine max [L.] Merr., var. Wye) plants. Continuous exposure of their rhizosphere to a pO2 of 0.06 atmospheres initially reduced nitrogenase activity by 37 to 45% with restoration of original activity in 4 to 24 hours and with no further change in tests up to 95 hours; continuous exposure to 0.02 atmosphere of O2 initially reduced nitrogenase activity 72%, with only partial recovery by 95 hours. Similar exposures to a pO2 of 0.32 atmospheres had little effect on N2[C2H2] fixation; a pO2 of 0.89 atmospheres initially reduced nitrogenase activity by 98% with restoration to only 14 to 24% of that of the ambient O2 controls by 95 hours. Re-exposure to ambient pO2 of plants adapted to nonambient pO2s reduced N2[C2H2] fixation to similar magnitudes as the reductions which occurred upon initial exposure to variant pO2 conditions, and a time period was required to readapt to ambient O2. It is concluded that the N2[C2H2]-fixing system of intact soybean plants is able to adapt to a wide range of external pO2s as probably occur in soil. We postulate that this occurs through an undefined mechanism which enables the nodule to maintain an internal pO2 optimal for nitrogenase activity.

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Adaptation of Nitrogen Fixation by Intact Soybean Nodules to Altered Rhizosphere pO2

Dry matter accumulation, nitrogen content and N(2) fixation rates of soybean (Glycine max [L.] Merr. cv. Wye) plants grown in chambers in which the aerial portion was exposed to a pO(2) of 5, 10, 21, or 30% and a pCO(2) of 300 mul CO(2)/l or a pO(2) of 21% and a pCO(2) of 1200 mul CO(2)/l during the complete growth cycle were measured. Total N(2)[C(2)H(2)] fixed was increased by CO(2)/O(2) ratios greater than those in air and was decreased by ratios smaller than those in air; the effects on N(2) fixation of decreased pO(2) or elevated pCO(2) were quantitatively similar during the period of vegetative growth. Decreased pO(2) produced a smaller increase then elevated pCO(2) during the reproductive period, presumably because of the decreased sink activity of the arrested reproductive growth under subambient pO(2). At a pO(2) of 5% and a pCO(2) of 300 mul CO(2)/l total N(2) fixed was increased 125% and per cent nitrogen content in the vegetative parts was increased relative to air while that in the seed was decreased. Dry matter production was increased and reproductive growth was arrested as previously reported for plants receiving only fertilizer nitrogen. At a pO(2) of 30% and a pCO(2) of 300 mul CO(2)/l total N(2) fixed was decreased 50% and per cent nitrogen content in the vegetative part was increased relative to air while that in the reproductive structures was unaffected. Dry matter production was similarly decreased in both vegetative and reproductive structures. These effects of altered pO(2) in the aerial part on N(2) fixation are consistent with the hypothesis that the amount of photosynthate available to the nodule may be the most significant primary factor limiting N(2) fixation while sink activity of the reproductive structures may be a secondary factor.

Effect of Altered pO2 in the Aerial Part of Soybean on Symbiotic N2 Fixation

The total metabolic cost of soybean (Glycine max L. Mer Clark) nodule nitrogen fixation was empirically separated into respiration associated with electron flow through nitrogenase and respiration associated with maintenance of nodule function.Rates of CO(2) evolution and H(2) evolution from intact, nodulated root systems under Ar:O(2) atmospheres decreased in parallel when plants were maintained in an extended dark period. While H(2) evolution approached zero after 36 hours of darkness at 22 degrees C, CO(2) evolution rate remained at 38 degrees of the rate measured in light. Of the remaining CO(2) evolution, 62% was estimated to originate from the nodules and represents a measure of nodule maintenance respiration. The nodule maintenance requirement was temperature dependent and was estimated at 79 and 137 micromoles CO(2) (per gram dry weight nodule) per hour at 22 degrees C and 30 degrees C, respectively.The cost of N(2) fixation in terms of CO(2) evolved per electron pair utilized by nitrogenase was estimated from the slope of H(2) evolution rate versus CO(2) evolution rate. The cost was 2 moles CO(2) evolved per mole H(2) evolved and was independent of temperature.In this symbiosis, nodule maintenance consumed 22% of total respiratory energy while the functioning of nitrogenase consumed a further 52%. The remaining respiratory energy was calculated to be associated with ammonia assimilation, transport of reduced N, and H(2) evolution.

Experimental determination of the respiration associated with soybean/rhizobium nitrogenase function, nodule maintenance, and total nodule nitrogen fixation.

IT is well recognized from short term experiments that the photosynthetic and vegetative growth rates of plants with high rates of photorespiration, the so-called C3 or inefficient plants, are elevated 50 to 100% by sub-atmospheric concentrations of O2, while those of plants with low rates of photorespiration, the so-called C4 or efficient plants, is essentially unaffected1,2. The long term effect of O2 concentrations on vegetative and reproductive growth of either C3 or C4 plants has not, however, been reported.

Ralph W. F. Hardy

Papers

Reproductive Growth and Dry Matter Production of Glycine max (L.) Merr. in Response to Oxygen Concentration.

Adaptation of Nitrogen Fixation by Intact Soybean Nodules to Altered Rhizosphere pO2

Effect of Altered pO2 in the Aerial Part of Soybean on Symbiotic N2 Fixation

Experimental determination of the respiration associated with soybean/rhizobium nitrogenase function, nodule maintenance, and total nodule nitrogen fixation.

Oxygen as a New Factor controlling Reproductive Growth