Interactive effects of soil nitrogen and atmospheric carbon dioxide on root/rhizosphere carbon dioxide efflux from loblolly and ponderosa pine seedlings

We measured CO₂ efflux from intact root/rhizosphere systems of 155 day old loblolly (Pinus taeda L.) and ponderosa (Pinus ponderosa Dougl. ex Laws.) pine seedlings in order to study the effects of elevated atmospheric CO₂ on the below-ground carbon balance of coniferous tree seedlings. Seedlings were grown in sterilized sand culture, watered daily with either 1, 3.5 or 7 mt M NH ₄ ⁺ , and maintained in an atmosphere of either 35 or 70 Pa CO₂. Carbon dioxide efflux (mol CO₂ plant^–1 s^–1) from the root/rhizosphere system of both species significantly increased when seedlings were grown in elevated CO₂, primarily due to large increases in root mass. Specific CO₂ efflux (mol CO₂ g root^–1 s^–1) responded to CO₂ only under conditions of adequate soil nitrogen availability (3.5 mt M). Under these conditions, CO₂ efflux rates from loblolly pine increased 70% from 0.0089 to 0.0151 mol g^–1 s^–1 with elevated CO₂ while ponderosa pine responded with a 59% decrease, from 0.0187 to 0.0077 mol g^–1 s^–1. Although below ground CO₂ efflux from seedlings grown in either sub-optimal (1 mt M) or supra-optimal (7 mt M) nitrogen availability did not respond to CO₂, there was a significant nitrogen treatment effect. Seedlings grown in supra-optimal soil nitrogen had significantly increased specific CO₂ efflux rates, and significantly lower total biomass compared to either of the other two nitrogen treatments. These results indicate that carbon losses from the root/rhizosphere systems are responsive to environmental resource availability, that the magnitude and direction of these responses are species dependent, and may lead to significantly different effects on whole plant carbon balance of these two forest tree species.     

The growth and nutrient uptake of winter wheat

Summary Measurements with an ion selective electrode under winter wheat and in adjacent fallow soil, from April to July 1976, showed that nitrate concentrations were high in the 0–25 cm zone and correspondingly lower at 50 cm, because of the extreme drying conditions. Maximum differences in nitrate concentrations between cropped and fallow soil occurred at Feekes' stages 6, 10, and 11.1 indicating periods of maximum uptake by the crop (cf Ref.⁴).Dry matter weight of wheat, sampled biweekly, was maximum 15 days before maturity. The foliage senesced and lost weight from Feekes' stage 10.1 onwards. Nutrient concentrations in the foliage decreased from Feekes' stage 4, but N, P and Mg concentrations in the ears increased during Feekes' stage 11. N, P and Mg accumulated in the ears at the expense of the foliage during stage 11, maximum uptake occurring at stages 11.3, 11.1 and 11.2 respectively. In contrast, K and Ca uptake ceased at stages 10.1 and 10.5 respectively and then both were lost from the foliage in heavy showers.Rates of N uptake and soil nitrate depletion correlated significantly, enabling N uptake to be deduced approximately from thesein situ soil nitrate measurements.     

The growth, yield and nutrient uptake of winter wheat following root-pruning

The effect of root-pruning on shoot growth was investigated in winter wheat growing in lysimeters. Removal of half of the root axes at the beginning of stem elongation reduced shoot dry matter, determined 1 month after pruning, by 13% and grain yield by 8%. Removal of either the seminal or nodal root system during tillering reduced shoot dry weight, measured during the growing season, by 7% and grain yield by 25%. Root-pruning had negligible, or only transient, effects on the concentration of nitrogen, phosphorus, potassium, magnesium and manganese in the shoots. The harvest index was not affected by root-pruning.     

Interactive effects of elevated carbon dioxide and growth temperature on photosynthesis in cotton leaves

Cotton (Gossypium hirsutum L., cv DPL 5415) plants were grown in naturally lit environment chambers at day/night temperature regimes of 26/18 (T-26/18), 31/23 (T-31/23) and 36/28 °C (T-36/28) and CO2 concentrations of 350 (C-350), 450 (C-450) and 700 L L-1 (C-700). Net photosynthesis rates, stomatal conductance, transpiration, RuBP carboxylase activity and the foliar contents of starch and sucrose were measured during different growth stages. Net CO2 assimilation rates increased with increasing CO2 and temperature regimes. The enhancement of photosynthesis was from 24 mol CO2 m-2 s-1 (with C-350 and T-26/18) to 41 mol m-2 s-1 (with C-700 and T-36/28). Stomatal conductance decreased with increasing CO2 while it increased up to T-31/23 and then declined. The interactive effects of CO2 and temperature resulted in a 30% decrease in transpiration. Although the leaves grown in elevated CO2 had high starch and sucrose concentrations, their content decreased with increasing temperature. Increasing temperature from T-26/18 to 36/28 increased RuBP carboxylase activity in the order of 121, 172 and 190 mol mg-1 chl h-1 at C-350, C-450 and C-700 respectively. Our data suggest that leaf photosynthesis in cotton benefited more from CO_2 enrichment at warm temperatures than at low growth temperature regimes.     

Effects of constant and fluctuating soil temperature on growth,development and nutrient uptake of maize seedlings

Summary The effect of constant and fluctuating soil temperature and two soil moisture regimes on the growth, development, transpiration and nutrient uptake by maize seedlings was studied in a greenhouse investigation. The constant root temperatures were maintained at 30, 34, 35, 36, 37, and 38°C for both 250 and 750 cm of soil moisture suctions. The fluctuating root temperature, for 250 cm of soil moisture suction only, of 30–35, 30–39, 30–40, 30–45 and 30–48°C were maintained to simulate the soil temperature regime under field conditions. The constant root temperature of 35°C and fluctuating temperature between 30–40°C significantly decreased the shoot and root growth and transpiration rate. On the average, there was 1.3 and 0.7 g decrease in fresh shoot weight and 0.36 and 0.30 g in fresh root weight per degree increase in root temperature for 250 and 750 soil moisture suction, respectively. In general, the effect of high soil moisture suction on maize seedlings was more severe when at high root temperature. The shoot and root concentration of N, P, and K decreased while that of B increased with increase in root temperature. The root concentration of Zn also decreased with increase in root temperature.     

Genotype and fungicide effects on late-season root growth of winter wheat

The aim of this work was to investigate differences among genotypes in post-anthesis root growth and distribution of modern UK winter wheat cultivars, and the effects of fungicide applications. Post-anthesis root growth of up to six cultivars of winter wheat (Triticum aestivum L.), given either one or three applications of fungicide, was studied in field experiments during two seasons. Total root mass remained unchanged between GS63 (anthesis) and GS85, but root length increased significantly from 14.7 to 31.4 km m⁻² in one season. Overall, there was no evidence for a decline in either root mass or length during grain filling. Root mass as a proportion of total plant mass was about 0.05 at GS85. There were significant differences among cultivars in root length and mass especially below 30 cm. Malacca had the smallest root length and Savannah the largest, and Shamrock had a significantly larger root system below 40 cm in both seasons. Fungicide applied at ear emergence had no significant effect on root mass in either season but increased root length (P<0.01) in the more disease-prone season. By maintaining a green canopy for longer, fungicide applied at flag leaf emergence may have resulted in delayed senescence of the root system and contributed to the post-anthesis maintenance of root mass and length. Section Editor: R. W. Bell     

The effects of increased atmospheric carbon dioxide and temperature on carbon partitioning, source-sink relations and respiration

Abstract. Herbaceous C₃ plants grown in elevated CO₂ show increases in carbon assimilation and carbohydrate accumulation (particularly starch) within source leaves. Although changes in the partitioning of biomass between root and shoot occur, the proportion of this extra assimilate made available for sink growth is not known. Root:shoot ratios tend to increase for CO₂-enriched herbaceous plants and decrease for CO₂-enriched trees. Root:shoot ratios for cereals tend to remain constant. In contrast, elevated temperatures decrease carbohydrate accumulation within source and sink regions of a plant and decrease root:shoot ratios. Allometric analysis of at least two species showing changes in root: shoot ratios due to elevated CO₂ show no alteration in the whole-plant partitioning of biomass. Little information is available for interactions between temperature and CO₂. Cold-adapted plants show little response to elevated levels of CO₂, with some species showing a decline in biomass accumulation. In general though, increasing temperature will increase sucrose synthesis, transport and utilization for CO₂-enriched plants and decrease carbohydrate accumulation within the leaf. Literature reports are discussed in relation to the hypothesis that sucrose is a major factor in the control of plant carbon partitioning. A model is presented in support.     

Effects of temperature on the development and growth of winter wheat roots

Winter wheat was sown on 2 dates with 3 levels of nitrogen fiertiliser (0, 50 and 200 kg N ha⁻¹) in one year and on 2 sites in a followign season. Shoot and root development and growth were measured between emergence and anthesis in the first season and emergence and 7 mainstem leaves in the second. Differences in temperature and light regime led to significant differences in shoot and root development and growth between sowing dates. A thermal time-scale, based on soil surface or air temperatures, with a base of 0°C, adequately described the production of mainstem leaves and nodal root axes over all treatments. Autumn applied nitrogen had little effect on development. Shoot growth and green area index increased exponentially with thermal time prior to spring nitrogen application and the completion of canopy development. Early-sown crops had larger root systems than late-sown crops prior to winter and this divergence was retained until anthesis. The relationship between root growth and thermal time was little better than with days after sowing and was not improved by either varying the site of temperature measurement or the base temperature used for calculation. Differences in soil texture and drainage, between sites, led to significant changes in root length distribution. Although spring applied nitrogen generally increased root length, its effects were inconsistent. There was a curvilinear relation between root length and the amount of photosynthetically active radiation (PAR) intercepted; this relation was unaffected by sowing date or nitrogen treatment. The amount of root produced per unit PAR decreased as the season progressed, reflecting the decrease in the proportion of total dry matter partitioned to the root system.     

Interactive effects of soil nutrient heterogeneity and belowground herbivory on the growth of plants with different root foraging traits

Aims

Plants with precise root foraging patterns can proliferate roots preferentially in nutrient-rich soil patches. When nutrients are distributed heterogeneously, this trait is often competitively advantageous in pot experiments but not field experiments. We hypothesized that this difference is due to belowground herbivory under field conditions.

Methods

We performed pot experiments using seedlings of Lolium perenne (a more precise root foraging species) and Plantago lanceolata (a less precise root foraging species). The experiment had a two-way factorial randomized block design, with nutrient distribution pattern (homogeneous or heterogeneous) and belowground herbivore (present or absent) as the two factors. Each pot contained one seedling of each species.

Results

With no herbivore present, plant biomass was smaller in the heterogeneous nutrient treatment than in the homogeneous treatment in P. lanceolata, but not in L. perenne. Under homogeneous nutrient distribution, plant biomass was lower in both species with a herbivore present than with no herbivore. Under heterogeneous nutrient distribution, biomass reduction due to herbivory occurred only in L. perenne.

Conclusions

Roots of the precise root foraging species were grazed more under the heterogeneous nutrient distribution, suggesting that the herbivore more efficiently foraged for roots in nutrient-rich soil patches.     

Cultivar differences in spatial root distribution during early growth in soil,and its relation to nutrient uptake - a study of wheat,onion and lettuce

Background and aims

A study was made to quantify early root development, soil exploitation and nutrient uptake in spring wheat, onion and lettuce, and their variation among cultivars. The goal was to study genetic variation in root traits making cultivars better adapted to organic production systems or other low-input systems.

Methods

Six cultivars of each species were grown in transparent tubes to allow direct observation of early root growth. The tubes were 0.3 m deep, and 0.24 m in diameter. By placing the plants close to the edge rather than at the centre of the tubes, we could quantify the spatial distribution of the root systems as well as the general root growth and nutrient uptake.

Results

Root growth of wheat and lettuce was faster than root growth of onion, and onion showed little capacity for horizontal root system development. Significant variation in early root growth and horizontal spread of the root system was found among cultivars of all three species. In general, cultivars with strong growth and high volume of soil exploitation showed higher average nutrient concentrations.

Conclusion

Early shoot growth, root growth and nutrient uptake are intrinsically linked, making it difficult to determine whether improved root growth was the primary cause of improved performance. However, we did find cultivars where the strong root growth and superior root distribution seemed to be the driver for improved overall growth.

     

Modelling the interacting effects of nutrient uptake, light capture and temperature on phytoplankton growth

A model of phytoplankton growth developed by analogy with chemicalkinetics (CR model) in Baird and Emsley (J. Plankton Res., 21,85–126, 1999) is explored further. The CR model parameterizesall biochemical reactions involved in phytoplankton growth byone parameter: the maximum growth rate. Phytoplankton growthrate is then calculated from an interaction of the maximum growthrate, and the physical limit to extracellular nutrient uptakerates and light capture. In this paper, the CR model was re-derived,with two corrections and a number of modifications to increaseits generality. During derivation, the model's behaviour wascompared with chemostat cultures at a variety of dilution rates,nutrient inputs and temperatures. Model output was then plottedagainst observations of a semi-continuous culture of Isochrysisgalbana. Finally, the CR model was used to predict the growthrate of phytoplankton communities extracted from two temperatelakes under varying nutrient, light and temperature regimes.The CR model explained 37% of the variability of phytoplanktongrowth rate in cultures at environmental conditions similarto those of the lakes, compared with 25% explained by a non-linearbest fit to 324 growth experiments. The following paper in thisissue develops the CR model further, using it to predict stablecarbon isotope fractionation.     

Variety specific relationships between effects of rhizobacteria on root exudation,growth and nutrient uptake of soybean

Aims

It has been increasingly recognized that only distal lower order roots turn over actively within the <2 mm fine root system of trees. This study aimed to estimate fine root production and turnover rate based on lower order fine roots and their relations to soil variables in mangroves.

Methods

We conducted sequential coring in five natural mangrove forests at Dongzhai Bay, China. Annual fine root production and turnover rate were calculated based on the seasonal variations of the biomass and necromass of lower order roots or the whole fine root system.

Results

Annual fine root production and turnover rate ranged between 571 and 2838 g m^?2 and 1.46–5.96 yr^?1, respectively, estimated with lower order roots, and they were increased by 0–30 % and reduced by 13–48 %, respectively, estimated with the whole fine root system. Annual fine root production was 1–3.5 times higher than aboveground litter production and was positively related to soil carbon, nitrogen and phosphorus concentrations. Fine root turnover rate was negatively related to soil salinity.

Conclusions

Mangrove fine root turnover plays a more important role than aboveground litter production in soil C accumulation. Sites with higher soil nutrients and lower salinity favor fine root production and turnover, and thus favor soil C accumulation.

     

Shoot and root growth and nutrients uptake of wheat as affected by soil layers

The effect of soil layering on the growth and nutrient content of wheat shoots and roots was studied. PVC containers (120 cm long and 25 cm inside diameter) were filled with layers of loam and loamy sand. Both roots and shoots dry weight increased as the thickness of loam layer increased. The root:shoot ratios decreased throughout the growing season. The N, P and K content of the shoots peaked at two weeks before anthesis, while shoot dry weight peaked at anthesis. The ranges of shoot content of N, P and K at anthesis for the different treatments were 6–25, 8–25 and 5–25% of the total plant nutrients, respectively. Late in the season the translocation rate of nutrients from the shoots to the seeds were in the following order N>P>K.     

The effect of rhizoctonia root disease and applied nitrogen on growth,nitrogen uptake and nutrient concentrations in spring wheat

Root disease caused by Rhizoctonia solani is a common problem of spring wheat in South Australia. There are reports that nitrogen applications can reduce the incidence and severity of the disease. A glasshouse trail in pots examined the effects of disease and of applied nitrogen on wheat growth, and evaluated the utility of the basal stem nitrate concentration in diagnosing deficiency in plants with and without root disease. Plants were harvested at the mid-tillering stage. Shoot growth was increased by applied nitrogen until a maximum yield was attained, after which additional N had no effect on shoot yield. Root growth, however, responded positively only to low levels of applied N, after which it declined, and in the highest N treatment root mass was less than in the plants without applied N. Root disease caused severe reductions in plant growth, and both root and shoot mass were affected similarly. Even though growth of diseased plants responded positively to applied nitrogen the response was less than that of disease-free plants. The critical concentration of basal stem nitrate-N did not appear to be affected by root disease, and was estimated at 1200 mg kg^-1, consistent with other glasshouse data. The basal stem nitrate-N concentration, either in fresh or dried tissue, appeared a better diagnostic tool of N stress than did total shoot N concentration or content, because of sharper definition of critical concentrations. Concentrations of other nutrients in shoot tissue were affected differentially by both applied nitrogen and root disease, but generally did not reach critical levels, although phosphorus and magnesium appeared deficient in very disease-stressed plants.     

Root dynamics in a semi-natural grassland in relation to atmospheric carbon dioxide enrichment,soil water and shoot biomass

Plant responses to increasing atmospheric CO₂ concentrations have been studied intensively. However, the effects of elevated CO₂ on root dynamics, which is important for global carbon budgets as well as for nutrient cycling in ecosystems, has received much less attention. We used minirhizotrons inside open-top chambers to study the effects of elevated atmospheric carbon dioxide concentration on root dynamics in a nutrient-poor semi-natural grassland in central Sweden. We conducted our investigation over three consecutive growing seasons during which three treatments were applied at the site: Elevated (≈ 700 μmol mol^-1) and ambient (≈ 360 μmol mol^-1) chamber levels of CO₂ and a control, without a chamber. During 1997, a summer with two dry periods, the elevated treatment compared with ambient had 25% greater mean root counts, 65% greater above-ground biomass and 15% greater soil moisture. The chambers seemed responsible for changes in root dynamics, whereas the elevated CO₂ treatment in general increased the absolute sum of root counts compared with the ambient chamber. In 1998, a wet growing season, there were no significant differences in shoot biomass or root dynamics and both chamber treatments had lower soil moisture than the control. We found that as seasonal dryness increased, the ratio of elevated – ambient shoot biomass production increased while the root to shoot ratio decreased. We conclude that this grasslands response to elevated CO₂ is dependent on seasonal weather conditions and that CO₂ enrichment will most significantly increase production in such a grassland when under water stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Influence of temperature on root development and phosphate influx in winter wheat grown at different P levels

Root development was studied in winter wheat ( Triticum aestivum L. cv Starke II) grown at 5,10, 15 and 20°C in nutrient solutions with phosphate concentrations of 10, 100 or 1000 μM . The plants were grown for 38 days (5 and 10°C), 19 days (15°C) or 14 days (20°C). At the end of the cultivation period the phosphate influx in the roots was determined with ³²P-phosphate. Root development (lateral and seminal roof length and number) was monitored throughout the cultivation period on the same individuals by repeated (approximately every second day) photocopying of the roots for measurements with digitizer and appropriate software. The 5°C treatment yielded no laterals, and the seminals were only slightly affected by the different phosphate treatments. The 10 μM phosphate treatment gave high root:shoot dry weight ratio, high average lateral root length and high specific root length [m root (g root fresh weight)^-1]. The 1000 μM phosphate treatment yielded the highest number of laterals per m seminal root, and usually also the highest absolute numbers. Phosphate influx decreased with increased P status of the roots. It is argued that phosphate influx is dependent on factors such as P status, root geometry and relative root extension rate.     

Relationship of photosynthetic acclimation to changes of Rubisco activity in field-grown winter wheat and barley during growth in elevated carbon dioxide

The responses of photosynthesis, Rubisco activity, Rubisco protein, leaf carbohydrates and total soluble protein to three carbon dioxide treatments were studied in winter wheat [Triticum aestivum (L.)] and barley [Hordeum vulgare (L.)]. Barley and wheat plants were grown in small field plots during 1995 and 1996 in clear, acrylic chambers (1.2–2.4 m²) and were provided with continuous carbon dioxide fertilization at concentrations of 350, 525 and 700 mol mol^–1. Photosynthetic rates of barley penultimate leaves and wheat flag leaves measured at growth carbon dioxide concentrations decreased with leaf age in all three CO₂ treatments during 1995 and 1996. Photosynthetic acclimation to elevated CO₂ was observed on seven of eight measurement dates for barley and ten of eleven measurement dates for wheat over both years. Initial Rubisco activity, total soluble protein and Rubisco protein in barley penultimate leaves and wheat flag leaves also decreased with leaf age. Total Rubisco activity was not used because of enzyme degradation. There was a significant CO₂ treatment effect on initial Rubisco activity, total soluble protein and Rubisco protein for wheat in 1995 and 1996 and for barley in 1995. Responses of barley penultimate leaf Rubisco activity and leaf protein concentrations to elevated carbon dioxide were nonsignificant in 1996. A significant CO₂ treatment effect also was detected when means of Rubisco activity, soluble protein and Rubisco protein for wheat flag leaves were combined over harvests and years. These three flag leaf parameters were not significantly different in the 350 and 525 mol mol^–1 CO₂ treatments but were decreased during growth in 700 mol mol^–1 CO₂ relative to the other two CO₂ treatments. Ratios of photosynthesis at 700 and 350 mol mol^–1 were compared to ratios of Rubisco activity at 700 and 350 mol mol^–1 using wheat flag leaf data from 1995 and 1996. Regression analysis of these data were linear [y = 0.586 + 1.103t x (r² = 0.432)] and were significant at P 0.05. This result indicated that photosynthetic acclimation was positively correlated with changes of initial Rubisco activity in wheat flag leaves in response to CO₂ enrichment. Effects of elevated CO₂ on wheat leaf proteins during 1995 and 1996 and on barley during 1995 were consistent with an acceleration of senescence.     

Effect of elevated atmospheric CO2 concentration on soil and root respiration in winter wheat by using a respiration partitioning chamber

Soil respiration in a cropland is the sum of heterotrophic (mainly microorganisms) and autotrophic (root) respiration. The contribution of both these types to soil respiration needs to be understood to evaluate the effects of environmental change on soil carbon cycling and sequestration. In this paper, the effects of free-air CO₂ enrichment (FACE) on hetero- and autotrophic respiration in a wheat field were differentiated and evaluated by a novel split-root growth and gas collection system. Elevated atmospheric pCO₂ of approximately 200 μmol mol⁻¹ above the ambient pCO₂ significantly increased soil respiration by 15.1 and 14.8% at high nitrogen (HN) and low nitrogen (LN) application rates, respectively. The effect of elevated atmospheric pCO₂ on root respiration was not consistent across the wheat growth stages. Elevated pCO₂ significantly increased and decreased root respiration at the booting-heading stage (middle stage) and the late-filling stage (late stage), respectively, in HN and LN treatments; however, no significant effect was found at the jointing stage (early stage). Thus, the effect of increased pCO₂ on cumulative root respiration for the entire wheat growing season was not significant. Cumulative root respiration accounted for approximately 25–30% of cumulative soil respiration in the entire wheat growing season. Consequently, cumulative microbial respiration (soil respiration minus root respiration) increased by 22.5 and 21.1% due to elevated pCO₂ in HN and LN, respectively. High nitrogen application significantly increased root respiration at the late stage under both elevated pCO₂ and ambient pCO₂; however, no significant effects were found on cumulative soil respiration, root respiration, and microbial respiration. These findings suggest that heterotrophic respiration, which is influenced by increased substrate supplies from the plant to the soil, is the key process to determine C emission from agro-ecosystems with regard to future scenarios of enriched pCO₂.     

Low growth temperature effects a differential inhibition of photosynthesis in spring and winter wheat

In vivo room temperature chlorophyll a fluorescence coupled with CO₂ and O₂ exchange was measured to determine photosynthetic limitation(s) for spring and winter wheat (Triticum aestivum L.) grown at cold-hardening temperatures (5°C/5°C, day/night). Plants of comparable physiological stage, but grown at nonhardening temperatures (20°C/16°C, day/night) were used in comparison. Winter wheat cultivars grown at 5°C had light-saturated rates of CO₂ exchange and apparent photon yields for CO₂ exchange and O₂ evolution that were equal to or greater than those of winter cultivars grown at 20°C. In contrast, spring wheat cultivars grown at 5°C showed 35% lower apparent photon yields for CO₂ exchange and 25% lower light-saturated rates of CO₂ exchange compared to 20°C grown controls. The lower CO₂ exchange capacity is not associated with a lower efficiency of photosystem II activity measured as either the apparent photon yield for O₂ evolution, the ratio of variable to maximal fluorescence, or the level of reduced primary quinone electron acceptor maintained at steady-state photosynthesis, and is most likely associated with carbon metabolism. The lower CO₂ exchange capacity of the spring cultivars developed following long-term exposure to low temperature and did not occur following over-night exposure of nonhardened plants to 5°C.