Lower Photosynthetic Nitrogen Use Efficiency Research Articles

Do Higher Resource Capture Ability and Utilization Efficiency Facilitate the Successful Invasion of Exotic Plant? A Case Study of <i>Alternanthera philoxeroides</i>

We tested the hypothesis that introduced populations may have higher resource capture ability and utilization efficiency than native ones of invasive plants. We compared ecophysiological traits including maximum photosynthetic rate (Pmax), apparent quantum yield (Q), specific leaf area (SLA), photosynthetic energy use efficiency (PEUE), photosynthetic nitrogen use efficiency (PNUE), water use efficiency (WUE), mass-based and area-based leaf construction cost (CCmass and CCarea), and mass-based and area-based leaf nitrogen concentration (Nmass and Narea) between native (Argentina) and introduced (USA) populations of two varieties (North Apa and South Apo) of Alternanthera philoxeroides under common garden conditions in China. For Apo and Apa, Pmax, Q, Nmass and WUE were not significantly different between native and introduced populations; introduced populations had significantly lower SLA and lower CCmass but significantly higher Narea and CCarea than native ones. For Apa, the introduced populations showed significantly lower PEUE and lower PNUE while for Apo, PEUE and PNUE were not significantly different between native and introduced populations. The results indicated that introduced populations of A. philoxeroides do not show higher resource capture ability and resource utilization efficiency than their native ones in the common garden experiment, suggesting that these traits may not necessarily contribute to successful invasion of invasive plants.

Photosynthetic nitrogen and water use efficiency of acacia and eucalypt seedlings as afforestation species

The ecophysiological traits of acacia and eucalypt are important in assessing their suitability for afforestation. We measured the gas-exchange rate, the leaf dry mass per area (LMA) and the leaf nitrogen content of two acacia and four eucalypt species. Relative to the eucalypts, the acacias had lower leaf net photosynthetic rate (PN), lower photosynthetic nitrogen-use efficiency (PNUE), higher water-use efficiency (WUE), higher LMA and higher leaf nitrogen per unit area (Narea). No clear differences were observed within or between genera in the maximum rate of carboxylation (Vcmax) or the maximum rate of electron transport (Jmax), although these parameters tended to be higher in eucalypts. PNUE and LMA were negatively correlated. We conclude that acacias with higher LMA do not allocate nitrogen efficiently to photosynthetic system, explaining why their PN and PNUE were lower than in eucalypts.

Plant biodiversity in a larch plantation from the view point of photosynthetic nitrogen use efficiency in northeast China

Forest floor of larch species often provides growth habitat for many kinds of understory species because of relatively sparse structure in a larch canopy. A rich flora of forest understory species may play an essential role in maintaining fertility of a larch stand. An attempt was made to evaluate photosynthetic nitrogen use efficiency (PNUE) of many understory and overstory species according to their Raunkiaer lifeform. By studying 72 perennial deciduous species in a larch plantation in northeast China, marked photosynthetic differences between phanerophytes (Ph) and other three lifeforms of chamaephytes (Ch), hemicryptophytes (He), and cryptophytes (Cr) were found, with marginal differences found among Ch, He, and Cr. Ph species had much lower PNUE, and much lower values of rate of nitrogen allocation to chlorophyll (Chl./N) and nitrogen allocation to carboxylation processes (V cmax/N) were concurrently observed in Ph compared with the other three lifeforms. Ph had much lower leaf nitrogen per unit of projection area (N area) and specific leaf area (SLA, cm2 g–1). At lower SLA, for Ph species the change of PNUE with SLA was small, but these changes became very large at higher SLA for Ch, He, and Cr species. Our findings indicate that leaf morphological change is important for clarifying photosynthesis differences among species with different lifeform.

Ontogenetic transition of leaf physiology and anatomy from seedlings to mature trees of a rain forest pioneer tree, Macaranga gigantea

In a field study, we compared anatomy and diurnal gas exchange and chlorophyll fluorescence in sunlit mature leaves of Macaranga gigantea (Reichb. f. and Zoll.) Muell. seedlings, saplings, an adult tree and suckers originating from stumps. We tested the hypothesis that the pattern of resource use shifts across various life stages with ontogenetic changes in leaf anatomy and physiology. Among leaves of different developmental stages, seedling leaves were the smallest and thinnest, whereas adult tree leaves were the largest and thickest, and the air space within the lamina was largest in seedling leaves and smallest in adult tree leaves. Photosynthetic nitrogen-use efficiency (PNUE) was higher in seedling and sapling leaves than in adult tree leaves. Mean PNUE in seedling leaves was 1.6 times that in adult tree leaves. Nevertheless, among the developmental stages, net photosynthetic rate (Pn) per unit leaf area was lowest in seedling leaves because they have the lowest nitrogen (N) content per unit leaf area. In situ water vapor stomatal conductance (g(s) at a given leaf-to-air vapor pressure deficit was highest in sapling leaves, suggesting that they have a high hydraulic efficiency per unit leaf area. Among developmental stages, intrinsic water-use efficiency (Pn/g(s)) and photochemical capacity of photosystem II were lowest in seedling leaves. Sapling leaves had the highest N concentration and Pn per unit dry mass and the highest g(s), indicating that the gradual transition from the seedling stage to the sapling stage is accompanied by an accumulation of N in plant bodies and the development of hydraulic systems to counteract unfavorable environmental stresses. The properties of adult tree leaves (low PNUE, high carbon:N ratio, small and dense cells and thick lamina) indicate that, during the transition from the sapling stage to the adult tree stage, the priority of resource use in leaves gradually shifts from enhancement of photosynthetic performance to defense against herbivory and mechanical damage. Leaf morphology and physiology were coordinated with the differences in resource use at each life stage.

Vertical, horizontal and azimuthal variations in leaf photosynthetic characteristics within a Fagus crenata crown in relation to light acclimation

An understanding of spatial variations in gas exchange parameters in relation to the light environment is crucial for modeling canopy photosynthesis. We measured vertical, horizontal and azimuthal (north and south) variations in photosynthetic capacity (i.e., the maximum rate of carboxylation: Vcmax), nitrogen content (N), leaf mass per area (LMA) and chlorophyll content (Chl) in relation to relative photosynthetic photon flux (rPPF) within a Fagus crenata Blume crown. The horizontal gradient of rPPF was similar in magnitude to the vertical gradient of rPPF from the upper to the lower crown. The rPPF in the north quadrant of the crown was slightly lower than in the south quadrant. Nitrogen content per area (Narea), LMA and Vcmax were strictly proportional to rPPF, irrespective of the vertical direction, horizontal direction and crown azimuth, whereas nitrogen content per dry mass, Chl per area and photosynthetic capacity per dry mass (Vm) were fairly constant. Statistical analyses separating vertical trends from horizontal and azimuthal trends indicated that, although horizontal and vertical light acclimation of leaf properties were similar, there were two significant azimuthal variations: (1) Vcmax was lower in north-facing leaves than in south-facing leaves for a given Narea, indicating low photosynthetic nitrogen-use efficiency (PNUE) of north-facing leaves; and (2) Vcmax was lower in north-facing leaves than in south-facing leaves for a given LMA, indicating low Vm of the north-facing leaves. With respect to the low PNUE of the north-facing leaves, there were no significant azimuthal variations in leaf CO2 conductance from the stomata to the carboxylation site. Biochemical analysis indicated that azimuthal variations in nitrogen allocation to ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and in nitrogen allocation between carboxylation (Rubisco and other Calvin cycle enzymes) and light harvesting machinery (Chl pigment-protein complexes) were not the main contributor to the difference in PNUE between north- and south-facing leaves. Lower specific activity of Rubisco may be responsible for the low PNUE of the north-facing leaves. Anatomical analysis indicated that not only high leaf density, which is compatible with a greater fraction of non-photosynthetic tissue, but also thick photosynthetic tissue contributed to the low Vm in the north-facing leaves. These azimuthal variations may need to be considered when modeling canopy photosynthesis based on the Narea-Vcmax or LMA-Vcmax relationship.

Interspecific difference in the photosynthesis?nitrogen relationship: patterns, physiological causes, and ecological importance

The photosynthesis-nitrogen relationship is significantly different among species. Photosynthetic capacity per unit leaf nitrogen, termed as photosynthetic nitrogen-use efficiency (PNUE), has been considered an important leaf trait to characterise species in relation to their leaf economics, physiology, and strategy. In this review, I discuss (1) relations between PNUE and species ecology, (2) physiological causes and (3) ecological implications of the interspecific difference in PNUE. Species with a high PNUE tend to have high growth rates and occur in disturbed or high productivity habitats, while those with a low PNUE occur in stressful or low productivity habitats. PNUE is an important leaf trait that correlates with other leaf traits, such as leaf mass per area (LMA) and leaf life span, irrespective of life form, phylogeny, and biomes. Various factors are involved in the interspecific difference. In particular, nitrogen allocation within leaves and the mesophyll conductance for CO(2) diffusion are important. To produce tough leaves, plants need to allocate more biomass and nitrogen to make thick cell walls, leading to a reduction in the mesophyll conductance and in nitrogen allocation to the photosynthetic apparatus. Allocation of biomass and nitrogen to cell walls may cause the negative relationship between PNUE and LMA. Since plants cannot maximise both PNUE and leaf toughness, there is a trade-off between photosynthesis and persistence, which enables the existence of species with various leaf characteristics on the earth.

Photosynthetic performance and resource utilization of two mangrove species coexisting in a hypersaline scrub forest.

In a hypersaline mangrove scrub forest in northern Florida, coexisting trees of Laguncularia racemosa and Avicennia germinans were either fertilized with nitrogen or phosphorus, or not fertilized (controls). We aimed to test whether nutrient additions differentially altered photosynthetic performance and resource utilization in these two species. In control trees, photosynthetic rates were higher in L. racemosa than A. germinans. However, leaf nitrogen concentrations were higher in A. germinans than L. racemosa. Avicennia germinans responded to fertilization with nitrogen by increasing leaf nitrogen concentrations and rates of photosynthesis such that they were equivalent to photosynthesis in L. racemosa. Laguncularia racemosa did not show a response to nitrogen additions. Neither species showed strong responses to phosphorus fertilization. Avicennia germinans had high photosynthetic water-use efficiency (photosynthesis/transpiration), but low photosynthetic nitrogen-use efficiency (photosynthesis/leaf nitrogen). In contrast, L. racemosa had comparatively low photosynthetic water use efficiency and high photosynthetic nitrogen use efficiency. Leaf level characteristics lead us to hypothesize that coexistence of A. germinans and L. racemosa should occur where nitrogen levels are low and salinity is moderate, or at least moderate for some period of the year.

Quercus species differ in water and nutrient characteristics in a resource-limited fall-line sandhill habitat.

We compared co-occurring mature Quercus laevis Walt. (turkey oak), Q. margaretta Ashe (sand post oak) and Q. incana Bartr. (bluejack oak) trees growing in resource-limited sandhill habitats of the southeastern United States for water and nutrient characteristics. The Quercus spp. differed in their distribution along soil water and nutrient gradients, and in their access to and use of water, even though the study year was wetter than average with no mid-season drought. Quercus laevis had the greatest access to soil water (least negative pre-dawn water potential, psi(pd)) and the most conservative water-use strategy based on its relatively low stomatal conductance (g(s)), high instantaneous water-use efficiency (WUE), least negative midday water potential (psy(md)) and high leaf specific hydraulic conductance (K(L)). Quercus margaretta had the least conservative water-use characteristics, exhibiting relatively high g(s), low instantaneous WUE, most negative psi(md), and low K(L). Quercus margaretta also had a low photosynthetic nitrogen-use efficiency (PNUE), but a high leaf phosphorus concentration. Quercus incana had the poorest access to soil water, but intermediate water-use characteristics and leaf nutrient characteristics more similar to those of Q. laevis. There were no species differences for photosynthesis (A), leaf nitrogen on an area basis, or seasonally integrated WUE (delta13C). Both A and g(s) were positively correlated for each species, but A and g(s) were generally not correlated with psi(pd), psi(md) or delta psi(pd-md). Although we found differences in resource use and resource status among these sandhill Quercus spp., the results are consistent with the interpretation that they are generally drought avoiders. Quercus laevis may have an advantage on xeric ridges because of its greater ability to access soil water and use it more conservatively compared with the other Quercus spp.

Optimal leaf area indices in C3 and C4 mono‐ and dicotyledonous species at low and high nitrogen availability

From an analytical model it was shown that for a given total amount of nitrogen in the canopy, there exists an optimal leaf area index (LAI), and therefore an optimal average leaf introgen content, at which canopy photosynthesis is maximal. If the LAI is increased above this optimum, increased light interception will not compensate for reduction in photosynthetic capacity of the canopy resulting from reduced leaf nitrogen contents. It was further derived from the model that the value of the optimal LAI increases with the photosynthetic nitrogen use efficiency (PNUE) and decreases with the canopy extinction coefficient for light (KL) and incident photon flux density (PFD) at the top of the canopy. These hypotheses were tested on dense stands of species with different photosynthetic modes and different architectures. A garden experiment was carried out with the C4 monocot sorghum (Sorghum bicolor [L.] Moensch cv. Pioneer), the C3 monocot rice (Oryza sativa L. cv. Araure 4), the C4 dicot amaranth (Amaranthus cruentus L. cv. K113) and the C3 dicot soybean (Glycine max [L.] Merr. cv. Williams) at two levels of nitrogen availability.The C4 species had higher PNUEs than the C3 species while the dicots formed stands with higher extinction coefficients for light and had lower PNUEs than the monocots. The C4 and monocot species were found to have formed more leaf area per unit leaf nitrogen (i.e., had lower leaf nitrogen contents) than the C3 and dicot species, respectively. These results indicate that the PNUE and the extinction coefficient for light are important factors determining the amount of leaf area produced per unit nitrogen as was predicted by the model.

Association of water use efficiency and nitrogen use efficiency with photosynthetic characteristics of two wheat cultivars

Abstract We examined if higher plant water use efficiency is related to a higher investment of nitrogen in components of the photosynthetic apparatus and if this is associated with a less efficient use of nitrogen for photosynthesis or for growth. We also assessed the influence of a reduced supply of water and nitrogen on these relationships. For this purpose we used two Triticum aestivum cultivars, differing in whole plant water use efficiency. Differences in leaf water use efficiency (A/E) between cultivars or treatments were associated with differences in Rubisco activity or rate of electron transport and, in general, with differences in plant water use efficiency. They were not invariably related to a low photosynthetic nitrogen use efficiency (PNUE). A high A/E was associated with a low PNUE, e.g. when we compared cultivars at high irradiance. In this case, a high A/E and a high leaf nitrogen concentration were associated with a low stornatal conductance and an equal rate of photosynthesis. A high A/E was not associated with a low PNUE when differences in A/E were mainly caused by higher rates of photosynthesis associated with higher nitrogen concentrations, e.g. when we compared cultivars at low irradiance and high water and nitrogen supply. An increase in the proportion of total leaf nitrogen associated with the photosynthetic apparatus at low soil-nitrogen increased PNUE, without affecting A/E in the opposite way. We conclude that whole plant nitrogen use efficiency was low for the cultivars with a high plant water use efficiency.

Photosynthetic capacity and nitrogen partitioning among species in the canopy of a herbaceous plant community.

Partitioning of nitrogen among species was determined in a stand of a tall herbaceous community. Total amount of nitrogen in the aboveground biomass was 261 mmol N m-2, of which 92% was in three dominant species (Phragmites, Calamagrostis and Carex) and the rest was in the other eight subordinate species. Higher nitrogen concentrations per unit leaf area (n L) with increasing photosynthetically active photon flux density (PPFD) were observed in all species except for three short species. The changes in n L within species were mainly explained by the different nitrogen concentrations per unit leaf mass, while the differences in n L between species were explained by the different SLM (leaf mass per unit leaf area). Photon absorption per unit leaf nitrogen (Φ N ) was determined for each species. If photosynthetic activity was proportional to photon absorption, Φ N should indicate in situ PNUE (photosynthetic nitrogen use efficiency). High Φ N of Calamagrostis (dominant) resulted from high photon absorption per unit leaf area (Φ area ), whereas high Φ N of Scutellaria (subordinate) resulted from low n L although its Φ area was low. Species with cylinder-like "leaves" (Juncus and Equisetum) had low Φ N , which resulted from their high n L. Light-saturated CO2 exchange rates per unit leaf area (CER) and per unit leaf nitrogen (potential PNUE) were determined in seven species. Species with high CER and high n L (Phragmites, Carex and Juncus) had low potential PNUE, while species with low CER and low n L showed high potential PNUE. NUE (ratio of dry mass production to nitrogen uptake) was approximated as a reciprocal of plant nitrogen concentration. In most species, three measures of nitrogen use efficiency (NUE, Φ N and potential PNUE) showed strong conformity. Nitrogen use efficiency was high in Calamagrostis and Scutellaria, intermediate in Phragmites and relatively low in Carex. Nitrogen use efficiency of subordinate species was as high as or even higher than that of dominant species, which suggests that growth is co-limited by light and nitrogen in the subordinate species.

Variation in Characters Related to Leaf Photosynthesis in Wild Bean Populations

Leaf characters related to photosynthesis in wild bean (Phaseolus vulgaris L.) populations were surveyed to identify genetically useful traits. Fifty wild bean accessions collected from Mexico to Argentina were evaluated at two contrasting field locations. Canonical analysis distinguished four groups of accessions, corresponding to collection sites in Mexico, Guatemala, Peru, and Argentina. Mexican accessions had high photosynthetic rates, low soluble protein contents, and the highest instantaneous photosynthetic nitrogen‐use efficiency (PSNUE). Guatemalan accessions had low photosynthetic rates, low N content, and low PSNUE. Peruvian accessions had high photosynthetic rates, few stomates, high N, high soluble protein, high chlorophyll content, and high PSNUE. Argentinian accessions had low photosynthetic rates, few adaxial stomates, low N content, and low PSNUE. It is proposed that palisade cell size influences leaf photosynthesis by affecting specific leaf weight and therefore N content per unit leaf area. Significant variation within and between origins was observed for adaxial stomatal density. Differences in PSNUE, adaxial stomatal density, and palisade cell volume between individual accessions varied by a factor of 3, 5, and 3.5, respectively. We conclude that genetic variation for leaf photosynthetic characters exists in wild bean germplasm.