Relative growth rate correlates negatively with pathogen resistance in radish: the role of plant chemistry

Plant growth rate has frequently been associated with herbivore defence: a large investment in quantitative defence compounds occurs at the expense of growth. We tested whether such a relationship also holds for growth rate and pathogen resistance. For 15 radish (Raphanus sativus L.) cultivars, we determined the potential growth rate and the resistance to fungal wilt disease caused by Fusarium oxysporum. We subsequently aimed to explain a putative negative relationship between growth rate and resistance based on plant chemical composition. Both growth rate and resistance level varied greatly among cultivars. Moreover, there was a strong negative correlation between growth rate and resistance, i.e. there are costs associated with a high resistance level. Roots of slow-growing, resistant cultivars have a higher biomass density. Using pyrolysis mass spectrometry. we part1y explained variation in both growth rate and resistance in terms of the same change in chemical composition. Leaves of slow-growing, resistant cultivars contained more cell wall material. Surprisingly, roots of slow-growing, highly resistant cultivars contained significantly less cell wall material, and more cytoplasmic elements (proteins). We speculate that this higher protein concentration is related to high construction and turn-over costs and high metabolic activity. The latter in turn is thought to be responsible for a rapid and adequate resistance reaction, in which phenols may be involved.     

Effects of global environmental change on carbon partitioning in vegetative plants of Triticum aestivum and closely related Aegilops species

The use of fossil fuel is predicted to cause an increase of the atmospheric CO₂ concentration, which will affect the global pattern of temperature and precipitation. It is therefore essential to incorporate effects of temperature and water supply on carbon partitioning of plants to predict effects of elevated [CO₂] on growth and yield of Triticum aestivum. Although earlier papers have emphasized that elevated [CO₂] favours investment of biomass in roots relative to that in leaves, it has now become clear that these are indirect effects, due to the more rapid depletion of nutrients in the root environment as a consequence of enhanced growth. Broadly generalized, the effect of temperature on biomass allocation in the vegetative stage is that the relative investment of biomass in roots is lowest at a certain optimum temperature and increases at both higher and lower temperatures. This is found not only when the temperature of the entire plant is varied, but also when only root temperature is changed whilst shoot temperature is kept constant. Effects of temperature on the allocation pattern can be explained largely by the effect of root temperature on the roots' capacity to transport water. Effects of a shortage in water supply on carbon partitioning are unambiguous: roots receive relatively more carbon. The pattern of biomass allocation in the vegetative stage and variation in water-use efficiency are prime factors determining a plant's potential for early growth and yield in different environments. In a comparison of a range of T. aestivum cultivars, a high water-use efficiency at the plant level correlates positively with a large investment in both leaf and root biomass, a low stomatal conductance and a large investment in photosynthetic capacity. We also present evidence that a lower investment of biomass in roots is not only associated with lower respiratory costs for root growth, but also with lower specific costs for ion uptake. We suggest the combination of a number of traits in future wheat cultivars, i.e. a high investment of biomass in leaves, which have a low stomatal conductance and a high photosynthetic capacity, and a low investment of biomass in roots, which have low respiratory costs. Such cultivars are considered highly appropriate in a future world, especially in the dryer regions. Although variation for the desired traits already exists among wheat cultivars, it is much larger among wild Aegilops species, which can readily be crossed with T. aestivum. Such wild relatives may be exploited to develop new wheat cultivars well-adapted to changed climatic conditions.     

Cyanide-Resistant Root Respiration and Tap Root Formation in Two Subspecies of Hypochaeris radicata

Root respiration of the tap root forming species Hypochaeris radicata L. was measured during tap root formation. A comparison was made of two subspecies: H. radicata L. ssp. radicata L., a subspecies from relatively rich soils, and H. radicata L. ssp. ericetorum Van Soest, a subspecies from poor acidic soils. Root respiration was high and to a large extent inhibited by hydroxamic acid (SHAM) before the start of the tap root formation, indicating a high activity of an alternative non-phosphorylative electron transport chain. The rate of root respiration was much lower and less sensitive to SHAM when a considerable tap root was present. However, root respiration was also cyanide-resistant when a tap root was present, indicating that the alternative pathway was still present. A decreased rate of root respiration coincided with an increase of the content of storage carbohydrates, mainly in the tap root. The level of reducing sugars was constant throughout the experimental period, and it was concluded that the activity of the alternative oxidative pathway was significant in oxidation of sugars that could not be utilized for purposes like energy production, the formation of intermediates for growth or for storage. Root respiration decreased after the formation of a tap root. This decrease could neither be attributed to a gradual disappearance of the alternative chain, nor to a decreased level of reducing sugars. No differences in respiratory metabolism between the two subspecies have been observed, suggesting that a high activity of the alternative oxidative pathway is not significant in adaptation of the present two subspecies to relatively nutrient-rich or poor soils.     

Effects of Drought on Partitioning of Nitrogen in Two Wheat Varieties Differing in Drought-tolerance

A model was constructed to describe the translocation and partitioningof nitrogen on the seventh day after anthesis for well-wateredand droughted plants of two wheat varieties (Triticum aestivumL. cv. Warigal and Condor). The glasshouse-grown plants weredetillered so that a simplified model could be derived for themain stem. A 9-d drought treatment was imposed just after anthesisand this coincided with the period of endosperm cell divisionin the grains. Warigal, which had a higher grain yield thanCondor under drought, absorbed up to 15-times more nitrogenand translocated 1.5-fold more nitrogen to the shoot via thexylem. In both varieties, nitrogen redistributed from vegetativeorgans accounted for more than 60 per cent in control and 70per cent in droughted plants of the nitrogen needed for eargrowth. The net loss of nitrogen increased by 4-3 per cent inthe leaves, but decreased by 60 per cent in the stem under drought.Stem and roots appeared to play an important role in the nitrogeneconomy of droughted plants: less nitrogen was translocateddirectly to the grains from the senescing leaves and 40–60per cent more nitrogen was translocated to the roots. Nearlyall the nitrogen reaching the roots in the phloem was reloadedinto the xylem stream and translocated back to the shoot. Thetransfer of nitrogen through the stem was reduced under droughtand this resulted in a constant C:N ratio of the grains whichmay be important in the regulation of endosperm cell division. Triticum aestivum L., wheat, drought, nitrogen, senescence, translocation     

The relationship between the relative growth rate and nitrogen economy of alpine and lowland Poa species

This study investigates the nitrogen economy of six altitudinally contrasting Poa species which differ in their relative growth rate (R). Two alpine (Poa fawcettiae and P. costiniana), one sub-alpine (P. alpina)and three temperate lowland species (P. pratensis, P. campressa and P. trivialis) were grown hydroponically under identical conditions in a growth room. The low R exhibited by the alpine species was associated with lower plant organic nitrogen concentration (n_p) and lower nitrogen productivity (Π_p, amount of biomass accumulation per mol organic nitrogen and time). The differences in Π_p between the alpine and lowland species did not appear to be due to differences in the carbon concentration or the proportion of total plant organic nitrogen allocated to the leaves, stems or roots. Variations in Π_P were also not due to variations in photosynthetic nitrogen use efficiency (Ψ_N, the rate of photosynthesis per unit organic leaf nitrogen) or shoot or root respiration rates per unit organic nitrogen (Λ_SH and Λ_R, respectively) per se. Rather, the lower Λ_p in the alpine species was probably due to a combination of small variations in several of the parameters (e.g. slightly lower Ψ_N, slightly higher Λ_SH and Λ_R, and slightly higher proportions of total plant organic nitrogen allocated to the roots). The alpine species exhibited lower organic acid and mineral concentrations. However, no differences in whole-plant construction costs (grams of glucose needed to synthesize one gram of biomass) were observed between She alpine and lowland Poa species. The lack of sub-stantial differences in Ψ_N between the alpine and lowland species contrasts with the large differences in Ψ_N between slow- and fast-growing lowland species that have been reported in the literature. The reasons for the unusually high Ψ_N values exhibited by the alpine Poa species are discussed.     

The physiological significance of cyanide-resistant respiration in higher plants

Abstract A brief survey of the biochemistry of the alternative oxidative pathway (‘cyanide-resistant respiration’) and its occurrence in vivo is given. Several hypotheses about the physiological significance of the alternative chain are discussed. These include a role in (1) heat production, (2) fruit ripening, (3) respiration of plants that contain high levels of cyanogenic glycosides, producing HCN upon wounding, (4) oxidation of NADH that is produced by various causes in excess of that required for ATP production, (5) ion uptake, and (6) osmoregulation. In intact roots of higher plants, the activity of the alternative pathway is more active when less carbohydrate is required for assimilation of N (NH+₄ NO-₃ or N₂) and is less active in those when carbohydrates are being stored in a storage organ or when the availability of photosynthate is reduced. An increase in carbohydrate requirement for osmoregulation is also correlated with decreased alternative chain activity. It is concluded that the alternative pathway in roots plays an important role in oxidation of sugars which are not required for carbon skeletons, energy production for growth and maintenance processes, osmoregulation or storage. However, the significance of this role may vary in different tissues and physiological states.     

Respiration of the Roots of Flood-Tolerant and Flood-Intolerant Senecio Species: Affinity for Oxygen and Resistance to Cyanide

The affinity of respiration for oxygen in the roots of six Senecio species studied was low compared with the affinity of cytochrome oxidase for oxygen. Half saturation values of approximately 22 μM oxygen were measured. Root respiration was to a large extent insensitive to cyanide in flood-tolerant as well as in flood-sensitive species. The evidence presented suggests that high activity of salicylhydroxamic acid (SHAM)-sensitive oxidase in Senecio roots was the basis for the low oxygen affinity and for the high cyanide-insensitivity of root respiration in the Senecio species. Methods are described to determine the in vivo activity of the SHAM-sensitive oxidase. It was estimated that it contributed 70% to the total root respiration. The presence of SHAM-sensitive oxidase activity could explain a higher efficiency of root growth respiration under a low oxygen tension if this alternate oxidase was inhibited at a low oxygen concentration in the root medium. However, the SHAM-sensitive oxidase was not specifically involved in either growth respiration or maintenance respiration. Its significance in regulation of the redox state of the cells is discussed.     

Efficiency and regulation of root respiration in a legume: Effects of the N source

A comparison was made of energy metabolism of nodulated N₂ fixing plants and non-nodulated NO₃-fed plants of Lupinus albus L. Growth, N-increment, root respiration (O₂ uptake and CO² production) and the contribution of a SHAM-sensitive oxidative pathway (the alternative pathway) in root respiration were measured. Both growth rate and the rate of N-increment were the same in both series of plants. The rate of root respiration, both O₂ uptake and CO₂ production, and the activity of the SHAM-sensitive pathway were higher in NO₃-fed plants than in N₂ fixing plants. The rate of ATP production in oxidative phosphorylation was computed also to be higher in NO₃-fed plants. It is concluded that both carbohydrate costings and ATP costings for synthesis + maintenance of root material were lower in N₂ fixing than in NO₃-fed plants. The respiratory quotient of root respiration was 1.6 in N₂-fixing plants and 1.4 in NO₃-fed plants. These values were slightly higher than the values calculated on the basis of CO₂ output due to N-assimilation and the experimental values of O₂ uptake, but showed the same trend: highest in N₂ fixing plants. Root respiration of NO₃-fed plants showed a diurnal pattern (both O₂ uptake, CO₂ production and the activity of the SHAM-sensitive pathway), whilst no diurnal variation in root respiration was found in N₂ fixing plants. However, C₂H₂ reduction did show a diurnal rhythm, which is suggested to be related to the diurnal variation in transpiration. Addition of NO₃ to N₂ fixing plants increased the rate of root respiration and the activity of the alternative pathway. This treatment did not decrease C₂H₂ reduction and H₂ evolution within 4 days. Withdrawal of NO₃-supply from NO₃-fed plants decreased the rate of root respiration but had no effect on the relative activity of the alternative pathway. It is suggested that the higher rate of root respiration and the higher activity of the SHAM-sensitive pathway in NO₃-fed plants is due to a larger supply of carbohydrates to the roots, partly due to a better photosynthetic performance of the shoots and partly due to a higher capacity of the roots to attract carbohydrates.     

Effects of nitrogen supply on the anatomy and chemical composition of leaves of four grass species belonging to the genus Poa, as determined by image-processing analysis and pyrolysis–mass spectrometry

Previous experiments have shown that the anatomy and chemical composition of leaves of inherently fast- and slow-growing grass species, grown at non-limiting nitrogen supply, differ systematically. The present experiment was carried out to investigate whether these differences persist when the plants are grown at an intermediate or a very low nitrogen supply. To this end, the inherently fast-growing Poa annua L. and Poa trivialis L., and the inherently slow-growing Poa compressa L. and Poa pratensis (L.) Schreb. were grown hydroponically at three levels of nitrate supply: at optimum (RGR_max) and at relative addition rates of 100 and 50 mmol N (mol N)^?1 d^?1 (RAR₁₀₀ and RAR₅₀), respectively. As expected, at the lowest N supply, the potentially fast-growing species grew at the same rate as the inherently slow-growing ones. Similarly, the differences in leaf area ratio (LAR, leaf area:total dry mass), specific leaf area (SLA, leaf arear:leaf dry mass) and leaf mass ratio (LMR, leaf dry mass:total dry mass) disappeared. Under optimal conditions, the fast-growing species differed from the slow-growing ones in that they had a higher N concentration. There were no significant differences in C concentration. With decreasing N supply, the total N concentration decreased and the differences between the species disappeared. The total C concentration increased for the fast-growing species and decreased for the slow-growing ones, i.e. the small, but insignificant, difference in C concentration between the species at RGR_max increased with decreasing N supply. The chemical composition of the leaves at low N supply, analysed in more detail by pyrolysis–mass spectrometry, showed an increase in the relative amounts of guaiacyl lignin, cellulose and hemicellulose, whereas those of syringyl lignin and protein decreased. The anatomy and morphology of the leaves of the four grass species differing in RGR_max were analysed by image-processing analysis. The proportion of the total volume occupied by mesophyll plus intercellular spaces and epidermis did not correlate with the amount of leaf mass per unit leaf area (specific leaf mass, SLM) at different N supply. The higher SLM at low N supply was caused partly by a high proportion of non-veinal sclerenchymatic cells per cross-section and partly by the smaller volume of epidermal cells. We conclude that the decrease in relative growth rate (and increase in SLM) at decreasing N supply is partly due to chemical and anatomical changes. The differences between the fast- and slow-growing grass species at an optimum nutrient supply diminished when plants were growing at a limiting nitrogen supply.