Critical N:P values: Predicting nutrient deficiencies in desert shrublands

Many terrestrial ecosystems are nutrient limited. Koerselman and Meuleman (1996) proposed critical foliar N:P values that could predict wetland nutrient deficiencies (N:P < 14, N limitation; N:P > 16, P limitation). Although critical N:P values have potential as ecological and diagnostic tools, species differences in N and P requirements must be tested. The objectives of our experiments were 1) to determine if two desert species share critical N:P values, and 2) to assess the N:P tool's applicability in a non-wetland system. We studied two common, co-occurring North American desert shrubs, Chrysothamnus nauseosus spp. consimilis and Sarcobatus vermiculatus. Based on leaf N and P concentrations, effective mycorrhizal status, and leaf N:P, we predicted Chrysothamnus would be N limited and Sarcobatus would be P limited. During the 2000 and 2001 growing seasons, juvenile shrubs received N, P, N+P, or control treatments. To assess interaction with water limitation, other shrubs were irrigated and fertilized in 2001 (N+IR, P+IR, N+P+IR, control IR). Contrary to the predicted N limitation in Chrysothamnus, stem growth was 1.7-fold greater in P+IR plants relative to IR controls, although without irrigation Chrysothamnus did not respond to P addition. Also, contrary to the predicted P limitation in Sarcobatus, stem growth was 2.7-fold and 1.5-fold greater in N and N+IR plants, respectively, relative to their controls. Leaf N was significantly higher in N-treated Sarcobatus, both with and without irrigation. Our study suggests species-specific critical N:P values and that the N:P tool does not effectively predict desert shrub nutrient limitations. Species adapted to low nutrient conditions may not respond to increased nutrient availability due to water and nutrient co-limitation, lack of plasticity, or lower tissue nutrient requirements.     

Nitrogen resorption from leaves under different growth irradiance in three deciduous woody species

Resorption of nitrogen (N) from senescing leaves is an important conservation mechanism that allows plants to use the same N repeatedly. We measured the extent of N resorption in plants co-occurring in a beech forest to examine the variability of N resorption, especially in relation to growth irradiance. Measurements were done in three deciduous woody species; one adult and several juvenile trees of Fagus crenata and several adult trees of Lindera umbellata and Magnolia salicifolia. N resorption efficiency (R_EFF; percentages of leaf N that is resorbed during leaf senescence) did not differ significantly among leaves under different growth irradiances in any species we studied. R_EFF was affected by the growth stage of the tree in F. crenata with the values being consistently lower in juvenile trees than in the adult tree. N resorption proficiency (R_PROF; N concentration of dead leaves) converged to a similar value in F. crenata juvenile trees and M. salicifolia, irrespective of the presenescent leaf N concentration that was affected by growth irradiance. Again, R_PROF was lower (i.e. absolute N concentration was higher) in juvenile trees than in the adult tree in F. crenata. These results suggest that the growth irradiance does not place a great impact on the extent of N resorption, but the growth stage of the tree is influential in some species. The difference between the adult and juvenile trees may be ascribed to the size of N sink tissues, which is likely to increase with plant age.     

Water relations and leaf chemistry of Chrysothamnus nauseosus ssp. consimilis (Asteraceae) and Sarcobatus vermiculatus (Chenopodiaceae)

At Mono Lake, California, we investigated field water relations, leaf and xylem chemistry, and gas exchange for two shrub species that commonly co-occur on marginally saline soils, and have similar life histories and rooting patterns. Both species had highest root length densities close to the surface and have large tap roots that probably reach ground water at 3.4-5.0 m on the study site. The species differed greatly in leaf water relations and leaf chemistry. Sarcobatus vermiculatus had a seasonal minimum predawn xylem pressure potential (ψ_pd) of -2.7 MPa and a midday potential (ψ_md) of -4.1 MPa. These were significantly lower than for Chrysothamnus nauseosus, which had a minimum ψ_pd of -1.0 MPa and ψ_md of -2.2 MPa. Sarcobatus had leaf Na of up to 9.1 % and K up to 2.7 % of dry mass, and these were significantly higher than for Chrysothamnus which had seasonal maxima of 0.4% leaf Na and 2.4 % leaf K. The molar ratios of leaf K/Na, Ca/Na, and Mg/Na were substantially lower for Sarcobatus than for Chrysothamnus. Xylem ionic contents indicated that both species excluded some Na at the root, but that Chrysothamnus was excluding much more than Sarcobatus. The higher Na content of Sarcobatus leaves was associated with greater leaf succulence, lower calculated osmotic potential, and lower xylem pressure potentials. Despite large differences in water relations and leaf chemistry, these species maintained similar diurnal patterns and rates of photosynthesis and stomatal conductance to water vapor diffusion. Sarcobatus ψ_pd may not reflect soil moisture availability due to root osmotic and hydraulic properties.     

Water potential and ionic effects on germination and seedling growth of two cold desert shrubs

We tested expectations that two desert shrubs would differ in germination and seedling relative growth rate (RGR) responses to Na and Ψ_s stress. The study species, Chrysothamnus nauseosus ssp. consimilis and Sarcobatus vermiculatus (hereafter referred to by genus), differ in their distribution along salinity gradients, with Chrysothamnus inhabiting only less saline areas. In growth chamber studies, declining Ψ_s (−0.82 to −2.71 MPa) inhibited germination of both species, and Chrysothamnus was less tolerant of Ψ_s stress than Sarcobatus. Germination fell below 10% for Chrysothamnus at −1.64 MPa (NaCl and PEG), and for Sarcobatus at −2.4 MPa PEG. Neither species exhibited ion toxicity. There was substantial ion enhancement for Sarcobatus in lower Ψ_s, allowing for 40% germination in −2.71 MPa NaCl. For seedling RGR, species were not different at −0.29 or −0.82 MPa (0 and 100 mmol/L NaCl, respectively), but Chrysothamnus RGR declined substantially at −1.3 MPa (200 mmol/L NaCl). The greater stress tolerance of Sarcobatus was not associated with a lower RGR under nonsaline conditions. Species differences in seed and seedling Ψ_s stress tolerance probably contribute to the restricted distribution of Chrysothamnus to less saline areas. The Na uptake of Sarcobatus seedlings enhances its ability to deal with declining Ψ_s and establish in more saline areas.     

EFFECT OF NITROGEN AND PHOSPHORUS SUPPLY ON GROWTH AND TISSUE COMPOSITION OF ULVA FENESTRATA AND ENTEROMORPHA INTESTINALIS (ULVALES,CHLOROPHYTA)1

The chlorophyte macroalgae Ulva fenestrata (Postels and Ruprecht) and Enteromorpha intestinalis (Linnaeus) Link. were grown under various nutrient regimes in indoor semi-continuous and batch cultures. Tissue nitrogen contents ranged from 1.3–5.4% N (dry wt), whereas tissue P ranged from 0.21–0.56% P (dry wt). Growth in low nitrogen medium resulted in N:P ratios of 5–8, whereas growth in high nitrogen medium resulted in N:P ratios of 21–44. For U. fenestrata, tissue N:P < 16 was indicative of N-limitation. Tissue N:P 16–24 was optimal for growth and tissue N:P > 24 was indicative of P-limitation. Growth of U. fenestrata was hyperbolically related to tissue N but linearly related to tissue P. Phosphorus-limited U. fenestrata maintained high levels of tissue N, but N-limited algae became depleted of P. For E. intestinalis, tissue N remained at maximum levels during P-limitation whereas tissue P decreased to about 85% of maximal levels during N-limitation. Growth rates for U. fenestrata decreased faster during P-limitation than during N-limitation. Simultaneously, tissue P was depleted faster than tissue N. Our results suggest that comparing tissue N and P of macroalage grown in batch cultures is useful for monitoring the nutritional status of macroalgae.     

生态恢复对马尾松叶片化学计量及氮磷转移的影响

为了解生态恢复对侵蚀红壤恢复的马尾松林叶片碳氮磷化学计量及氮磷养分转移的影响,在福建省长汀县河田镇典型侵蚀红壤区选取恢复13、30、33a的马尾松林为研究对象,并以未治理侵蚀地(CK1)和次生林(CK2)分别作为恢复前和恢复后的对照,通过测定马尾松叶片的碳、氮、磷含量,计算其计量比,内稳性指数和氮磷转移率,分析了侵蚀红壤生长的马尾松养分限制与养分转移的关系。结果表明:在侵蚀红壤恢复过程中,马尾松1年龄叶片C、N、P含量及1年龄叶片C∶N、C∶P、N∶P变化较小,这与马尾松较高的内稳性有关(N和P内稳性指数分别为7.57和3.89)。所有实验地马尾松1年龄叶片N∶P处于11.0—13.4之间,表明马尾松的生长受N、P共同限制,其中马尾松叶片N转移率显著低于P转移率,这与生态恢复过程中马尾松养分利用效率、生长需求以及土壤养分供应状况有关。1年龄叶片C∶N、C∶P分别与马尾松N、P转移率成负相关关系,当马尾松叶片C∶N、C∶P较低时,表明N、P利用效率较低,叶片衰老时更多的N和P被转移利用;反之,则N、P利用效率较高,转移率低。同时,C∶N、C∶P分别与树高、胸径成显著负相关关系,即马尾松生长对N、P的需求同样会影响化学计量比的变化,从而影响养分转移。虽然侵蚀地生态恢复过程中土壤N、P含量增加,但仍较贫瘠,不足以满足马尾松的生长,马尾松养分转移率较高,因此,为了提高侵蚀地恢复的马尾松林的生产力,建议下一步恢复措施中适当施加N肥和P肥。该研究将侵蚀红壤不同生态恢复年限的马尾松叶片C、N、P化学计量及养分转移结合,有助于全面、系统地揭示生态恢复过程马尾松林的养分循环,对指导侵蚀红壤恢复和提高马尾松生产力具有重要意义。     

Plant N capture from pulses: effects of pulse size,growth rate,and other soil resources

In arid ecosystems, the ability to rapidly capture nitrogen (N) from brief pulses is expected to influence plant growth, survival, and competitive ability. Theory and data suggest that N capture from pulses should depend on plant growth rate and availability of other limiting resources. Theory also predicts trade-offs in plant stress tolerance and ability to capture N from different size pulses. We injected K¹⁵NO₃, to simulate small and large N pulses at three different times during the growing season into soil around the co-dominant Great Basin species Sarcobatus vermiculatus, Chrysothamnus nauseosus ssp. consimilis, and Distichlis spicata. Soils were amended with water and P in a partial factorial design. As predicted, all study species showed a comparable decline in N capture from large pulses through the season as growth rates slowed. Surprisingly, however, water and P availability differentially influenced the ability of these species to capture N from pulses. Distichlis N capture increased up to tenfold with water addition while Chrysothamnus N capture increased up to threefold with P addition. Sarcobatus N capture was not affected by water or P availability. Opposite to our prediction, Sarcobatus, the most stress tolerant species, captured less N from small pulses but more N from large pulses relative to the other species. These observations suggest that variation in N pulse timing and size can interact with variable soil water and P supply to determine how N is partitioned among co-existing Great Basin species.     

Nitrogen resorption from senescing leaves of three salt marsh plant species

Seasonalvariation in leaf nitrogen of mature green and senescent leaves and nitrogenresorption efficiency in three plants (Spartina maritima, Halimioneportulacoides and Arthrocnemum perenne) of aTagus estuary salt marsh are reported. Total nitrogen concentrations in greenand senescent leaves were higher during winter (December and March). Soilinorganic nitrogen availability showed an opposite pattern with higherconcentrations during summer (June and September) when total leaf biomass washigher. Nitrogen resorption efficiency ranged between 31 and 76% andH. portulacoides was the plant that better minimizednitrogen loss by this process. Nitrogen resorption occurred mainly from thesoluble protein pool, although other fractions must have been broken down duringthe resorption process. No significant seasonal variation in nitrogen resorptionefficiency and no relation to leaf total nitrogen or soil nitrogen availabilitywere found. This suggests that the efficiency of the resorption process is notdetermined by the plant nitrogen status nor by the availability of the nutrientin the soil. Nevertheless, resorption from senescing leaves may play animportant role in the nitrogen dynamics of salt marsh plants and reduce thenitrogen requirements for plant growth.     

不同林龄短枝木麻黄小枝单宁含量及养分再吸收动态

通过对福建省惠安县不同林龄短枝木麻黄人工林不同发育阶段小枝单宁含量及氮磷再吸收率的研究,探讨了短枝木麻黄林分发育过程中的营养保存策略。结果表明,幼龄林(5年生)成熟小枝中的总酚(TP)、可溶性缩合单宁(ECT)总缩合单宁(TCT)含量及蛋白质结合能力(PPC)显著高于成熟林(21年生)和衰老林(38年生)。随着林分发育,成熟小枝中N含量显著升高,而P含量呈降低趋势。不同发育阶段林分成熟小枝中N:P比均大于20,且随着林龄的增加而升高。磷再吸收率(PRE)显著高于氮再吸收率(NRE),均以成熟林分最高。这表明短枝木麻黄小枝单宁含量与养分再吸收受林龄影响,其养分保存机制会随着林分生长发育的变化而进行调节。     

Nutrient resorption patterns of plant functional groups in a tropical savanna: variation and functional significance

Green and senesced leaf nitrogen (N) and phosphorus (P) concentrations of different plant functional groups in savanna communities of Kruger National Park, South Africa were analyzed to determine if nutrient resorption was regulated by plant nutritional status and foliar N:P ratios. The N and P concentrations in green leaves and the N concentrations in senesced leaves differed significantly between the dominant plant functional groups in these savannas: fine-leaved trees, broad-leaved trees and grasses. However, all three functional groups reduced P to comparable and very low levels in senesced leaves, suggesting that P was tightly conserved in this tropical semi-arid savanna ecosystem. Across all functional groups, there was evidence for nutritional control of resorption in this system, with both N and P resorption efficiencies decreasing as green leaf nutrient concentrations increased. However, specific patterns of resorption and the functional relationships between nutrient concentrations in green and senesced leaves varied by nutrient and plant functional group. Functional relationships between N concentrations in green and senesced leaves were indistinguishable between the dominant groups, suggesting that variation in N resorption efficiency was largely the result of inter-life form differences in green leaf N concentrations. In contrast, observed differences in P resorption efficiencies between life forms appear to be the result of both differences in green leaf P concentrations as well as inherent differences between life forms in the fraction of green leaf P resorbed from senescing leaves. Our results indicate that foliar N:P ratios are poor predictors of resorption efficiency in this ecosystem, in contrast to N and P resorption proficiencies, which are more responsive to foliar N:P ratios.     

Nitrogen resorption from needles of Pinus thunbergii Parl. growing along a topographic gradient of soil nutrient availability

To examine the relative importance of nutrient resorption in increasing the nutrient-use efficiency of Pinus thunbergii Parl., we investigated the nitrogen contents of green and senescent needles of P. thunbergii trees growing at five positions along a slope (LS, lower slope; TR, transitional site; MS, middle slope; US, upper slope; RG, ridge) and found that soil nitrogen availability tended to decrease upslope. Nitrogen concentration in green and senescent needles decreased upslope. Nitrogen resorption efficiencies (percentage change in nitrogen content between green and senescent needles) increased upslope from 43 to 77% with decreasing soil nitrogen availability. Nitrogen resorption efficiency was related to green needle dry mass per unit length, but there was no clear correlation between nitrogen resorption efficiency and nitrogen content in green needles. We concluded that the increase in nitrogen resorption efficiency of P. thunbergii enhanced the nitrogen-use efficiency as a response to the low nitrogen availability.     

Nutrient resorption response to fire and nitrogen addition in a semi-arid grassland

Fire and nitrogen (N) addition, both widely used grassland restoration strategies, strongly influence community composition and ecosystem functioning. However, little is known about their effects on plant nutrient resorption from senescing leaves, especially in semi-arid ecosystems. We evaluated the effects of fire, N addition (5.25 g N m⁻² yr⁻¹) and their potential interactions on nutrient resorption in five plant species in a semi-arid grassland in northern China. Foliar nutrient concentrations and resorption proficiencies and efficiencies varied substantially among species and functional groups. Fire increased green leaf N concentration ([N]g) and decreased N resorption proficiency (N RP), P resorption proficiency (P RP) and P resorption efficiency (P RE). N addition led to higher [N]g and lower N resorption, whereas it did not affect P related responses. There was no interaction between fire and N addition to affect all response variables except for green leaf P concentration ([P]g). These results suggest that fire and N addition can influence ecosystem nutrient cycling directly by changing resorption patterns and litter quality. Given the substantial interspecific variations in nutrient content and resorption and the potentially changing community composition, both fire and N addition may have indirect impacts on ecosystem nutrient cycling in this semi-arid grassland.     

Nitrogen addition increases fecundity in the desert shrub Sarcobatus vermiculatus

Nutrients, in addition to water, limit desert primary productivity, but nutrient limitations to fecundity and seed quality in desert ecosystems have received little attention. Reduced seed production and quality may affect recruitment, population, and community processes. At the Mono Basin, CA, USA where the alkaline, sandy soil has very low availability of N, P, and most other nutrients, seed production, recruitment, and dominance of the desert shrub Sarcobatus vermiculatus decrease over a dune successional sequence. Concurrently, Sarcobatus leaf N, P, and Ca/Mg ratio decline from early to later successional dunes. At two later successional dune sites, we fertilized adult Sarcobatus shrubs for 2 years and determined which nutrient(s) limited growth, seed production, and seed quality. We also tested whether nutrient addition at these older sites made these fitness-related variables equivalent to a younger, high-fecundity site. Nitrogen addition, alone, increased Sarcobatus leaf N, growth, and seed production per shoot module. Any treatment including P, Ca, Mg, or micronutrients but not N had an insignificant effect on growth and fecundity. Nitrogen addition also increased filled seed weight, a predictor of potential seedling survival, at one of the sites. Nitrogen-limited seed production and seed mass may reduce Sarcobatus fitness and contribute to the observed successional changes in plant community composition in this alkaline desert ecosystem.     

氮添加对黄河三角洲滨海湿地芦苇养分再吸收效率的影响

大气氮沉降增加能改变土壤养分可利用性,影响滨海湿地植物的养分再吸收。目前研究多关注氮沉降量对养分再吸收过程的影响,且研究集中于叶片,鲜有研究区分不同形态氮素对植物不同器官养分再吸收过程的影响。通过两年的野外控制实验,研究硝态氮、铵态氮添加对黄河三角洲滨海湿地芦苇(Phragmites australis)叶、茎养分再吸收效率的影响。结果表明：两类氮添加均显著增加叶、茎的氮、磷含量(P<0.001),增幅达32.74%—43.22%(氮)、30.91%—36.51%(磷)。叶片氮的再吸收效率为54.14%—67.66%,茎氮的再吸收效率为50.60%—62.85%。叶片磷的再吸收效率为56.80%—70.38%,茎磷的再吸收效率为77.43%—84.95%。两类氮添加均显著降低氮、磷的再吸收效率(P<0.001),但两类氮添加处理下的养分再吸收效率无差异。叶、茎氮的再吸收效率无差异,但茎磷的再吸收效率明显高于叶(P<0.01)。总之,氮添加降低芦苇对氮、磷的再吸收效率,且茎对养分的再吸收也具有不可忽略的贡献。     

Nutrient content in Quercus ilex canopies: Seasonal and spatial variation within a catchment

Spatial and temporal changes in canopy nutrient content were studied in 1988 and 1989 in a Mediterranean Quercus ilex ssp. ilex L. forest in north-eastern Spain. Sampling was conducted in parallel at two sites which represent endpoints along a slope gradient within a small catchment (ridge top at 975 m and valley bottom at 700 m). Deeper soils resulted in significantly higher N and P concentrations, and N content on a leaf area basis at the valley bottom site. In contrast, K concentration in leaves was significantly higher at the ridge top site, where soil K concentration was also higher. At both sites, N and P content on a leaf area basis was highest at the top of the canopy, where leaf area is highest. N resorption efficiency decreased from top to bottom of the canopy. Results suggested a minor role of shaded leaves as nutrient storage sites. Lower P resorption efficiency was found at the ridge top site. Seasonal changes of P and N concentration on a leaf area basis suggest P replenishment, and to a lesser degree N, during periods of lower growth activity due to low temperatures, but coinciding with higher water availability (autumn-early spring period). Thus, N and P resorption from the remaining foliage in the canopy took place, and to a larger degree at the valley bottom site, coinciding with a slightly higher leaf area index and productivity at this site.     

闽楠林土壤-凋落物-叶片碳氮磷生态化学计量对人为干扰的响应

干扰是影响森林生态系统稳定性和功能的重要因子,干扰程度直接影响天然林的生长进而影响其生态系统能量流动和养分循环过程,为此开展干扰对天然林生态系统影响研究,对于揭示干扰对天然林生态系统养分平衡特征机制具有重要意义。以福建两种人为干扰模式下(重度干扰和轻度干扰)闽楠林为研究对象,通过分析“土壤-凋落物-叶片”三个组分化学计量特征,结合养分利用效率、养分再吸收效率、内稳态理论解析干扰对闽楠林养分资源利用策略和生态适应。结果表明：(1)两种干扰模式下,叶片C、N、P含量均显著高于其土壤和凋落物,且三个组分中N和P含量均表现出重度干扰显著高于轻度干扰,但三个组分C/N、C/P和N/P呈现轻度干扰显著高于其重度干扰。(2)闽楠林叶片N、P养分利用效率表现出：重度干扰<轻度干扰,但P再吸收效率则是重度干扰高于轻度干扰,且两种干扰模式下P养分利用效率和再吸收效率显著高于N。(3)随干扰强度的增加,闽楠林叶片N呈现内稳态弱,而叶片P的内稳态强以适应低P环境。(4)凋落物与叶片两组分P、C/P、N/P存在显著正相关关系,土壤C/N分别与叶片P、C/P、N/P以及凋落物P、C/N和C/P存在显著相关关...     

Nutrient resorption of two evergreen shrubs in response to long-term fertilization in a bog

Plant resorption of multiple nutrients during leaf senescence has been established but stoichiometric changes among N, P and K during resorption and after fertilization are poorly understood. We anticipated that increased N supply would lead to further P limitation or co-limitation with N or K [i.e. P-(co)limitation], decrease N resorption and increase P and K resorption, while P and K addition would decrease P and K resorption and increase N resorption. Furthermore, Ca would accumulate while Mg would be resorbed during leaf senescence, irrespective of fertilization. We investigated the effect of N, P and K addition on resorption in two evergreen shrubs (Chamaedaphne calyculata and Rhododendron groenlandicum) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada. In general, N addition caused further P-(co)limitation, increased P and K resorption efficiency but did not affect N resorption. P and K addition did not shift the system to N limitation and affect K resorption, but reduced P resorption proficiency. C. calyculata resorbed both Ca and Mg while R. groenlandicum resorbed neither. C. calyculata showed a higher resorption than R. groenlandicum, suggesting it is better adapted to nutrient deficiency than R. groenlandicum. Resorption during leaf senescence decreased N:P, N:K and K:P ratios. The limited response of N and K and the response of P resorption to fertilization reflect the stoichiometric coupling of nutrient cycling, which varies among the two shrub species; changes in species composition may affect nutrient cycling in bogs.     

Effects of different levels of N- and P-deficiency on cell yield,okadaic acid,DTX-1, protein and carbohydrate dynamics in the benthic dinoflagellate Prorocentrum lima

Prorocentrum lima (Ehrenberg) Dodge is a cosmopolitan epiphytic dinoflagellate that produces biotoxins which are causative of diarrhetic shellfish poisoning (DPS). Here we report on effects of several nitrogen (N) and phosphorous (P) limited conditions on cell yield, okadaic acid (OA) and dinophysistoxin-1 (DTX-1) contents synoptically with cell carbohydrate, exopolysaccharide (EPS) and cell protein concentrations in a P. lima strain isolated from the Sacca di Goro lagoon (Northern Adriatic Sea). Batch culture experiments were set to assess changes induced by four nitrogen-limited levels (1/3-N, 1/10-N, 1/20-N, and 1/50-N) and four phosphorus-limited levels (1/3-P, 1/10-P, 1/20-P, and 1/50-P) with respect to control nutrient conditions (f/2 medium; NO₃⁻ and PO₄³⁻ concentrations: 883 and 36.3 μM, respectively; N/P ratio: 24). Low nutrients availability determined lower cell yields starting from 1/10-N and 1/3-P levels and the pattern observed was dependent on nutrient dynamics, as shown by N and P analyses performed in culture media during growth. Final cell yield decreased significantly up to 4.7- and 5.6-fold under 1/50-N and 1/50-P-limited levels with respect to control values, while cell volume increased with respect to control (up to 30% and 35% for N- and P-experiment, respectively). On overall, OA concentration ranged from 6.69 to 15.80 pg cell⁻¹, while DTX-1 ranged from 0.12 and 0.39 pg cell⁻¹ resulting in unusual high OA/DTX-1 ratios. The study indicates that protein, carbohydrate, EPS, and toxin concentrations displayed remarkable different patterns under the two kinds of nutrient deficiencies. The main differences can be summarised as: (i) significant decrease of cell protein concentration (up to 2-fold) under N-limitation, conversely no significant changes in protein concentration under P-limitation; (ii) significant increase of cell carbohydrate (up to 2.8-fold and 3.4-fold for N- and P-limitation, respectively) and cell OA amount (up to 1.9-fold and 2.3-fold, N- and P-limitation, respectively) under both N- and P-limitations, however different level-deficiency dependent patterns were displayed under the two nutrient conditions; (iii) significant increase of EPS concentration (up to 6.50-fold) under P-limitation, conversely no significant changes in EPS concentration under N-limitation. Data presented here indicate that P. lima adopts different eco-physiological strategies to face N-limitation or P-limitation. This study provides the first evidence for an increase in EPS production by benthic dinoflagellates under P-limited conditions; the ecological significance of this increase is discussed.     

Leaf litter decomposition and nutrient dynamics in a subtropical forest after typhoon disturbance

Decomposition of typhoon-generated and normal leaf litter and their release patterns for eight nutrient elements were investigated over 3 yr using the litterbag technique in a subtropical evergreen broad-leaved forest on Okinawa Island, Japan. Two common tree species, Castanopsis sieboldii and Schima wallichii, representative of the vegetation and differing in their foliar traits, were selected. The elements analyzed were N, P, K, Ca, Mg, Na, Al, Fe and Mn. Dry mass loss at the end of study varied in the order: typhoon green leaves > typhoon yellow leaves > normal leaves falling for both species. For the same litter type, Schima decomposed faster than Castanopsis. Dry mass remaining after 2 yr of decomposition was positively correlated with initial C:N and C:P ratios. There was a wide range in patterns of nutrient concentration, from a net accumulation to a rapid loss in decomposition. Leaf litter generated by typhoons decomposed more rapidly than did the normal litter, with rapid losses for N and P. Analysis of initial quality for the different litter types showed that the C:P ratios were extremely high (range 896 – 2467) but the P:N ratios were < 0.05 (range 0.02 – 0.04), indicating a likely P-limitation for this forest. On average 32% less N and 60% less P was retranslocated from the typhoon-generated green leaves than from the normal litter for the two species, Castanopsis and Schima. An estimated 2.13 g m^–2 yr^–1 more N and 0.07 g m^–2 yr^–1 more P was transferred to the soil as result of typhoon disturbances, which were as high as 52% of N and 74% of P inputted from leaf litter annually in a normal year. Typhoon-driven maintenance of rapid P cycling appears to be an important mechanism by which growth of this Okinawan subtropical forest is maintained.     

Changes of Photosynthetic Characteristics in Relation to Leaf Senescence in Two Maize Hybrids with Different Senescent Appearance

A field experiment was conducted to investigate the changes in chlorophyll (Chl) and nitrogen (N) contents, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and phosphoenolpyruvate carboxylase (PEPC) contents and PEPC activity, and the photon-saturated net photosynthetic rate (P _Nsat), and their relationships with leaf senescence in two maize hybrids with different senescent appearance. One stay-green (cv. P3845) and one earlier senescent (cv. Hokkou 55) hybrid were used in this study, and we found that Chl and N contents and the P _Nsat in individual leaves of P3845 were greater than those in corresponding leaves of Hokkou 55 at the successive growth stages. In addition, larger contents of RuBPCO and PEPC, and a greater activity of PEPC were observed in P3845. Due to the lower rates of decrease of Chl, RuBPCO, and PEPC amounts per unit of N, and the lower net C translocation rate per unit of N in the stay-green hybrid, leaf senescence was delayed in comparison to the earlier senescent hybrid.