Phosphorus limitation and competitive capacities of Pinus halepensis and Quercus ilex subsp. rotundifolia on different soils

Aleppo pine (Pinus halepensis) and the evergreen holm oak (Quercus ilex) dominate forest areas of the Mediterranean Basin. Both species regenerate abundantly after fires: pine through seedlings and holm oak through resprouts. Cumulative nutrient losses caused by frequent fires may have decreased soil nutrient availability in such areas. To assess the role of nitrogen and phosphorus as limiting factors for growth of these species during post-fire recovery, a field fertilisation and competition experiment was conducted in a 5-year post-fire shrubland on calcareous soil, where naturally-regenerated saplings of Aleppo pine and resprouts of interior holm oak (Quercus ilex subsp. rotundifolia) coexist. Three years after fertilisation, relative basal area increment was 56% greater in pines fertilised with 250 kg P ha^–1 than in non fertilised ones. N fertilisation had small or no effects. Interactions between N and P fertilisation were not observed. Growth of Aleppo pine only increased with P fertilisation when neighbours were removed. Hence, the negative effect of neighbours on growth was greater when P availability was enhanced by fertilisation. In contrast, holm oak was able to grow more (110%) in response to increased P supply even without neighbour removal. A common garden experiment was then conducted with potted seedlings to investigate whether the suggested higher competitive capacity of holm oak for P held under a range of P amendments on different soils and competitive situations. P fertilisation increased seedling biomass yield of both species. When P availability increased, a negative effect of neighbours on growth was observed for holm oak and in 70 a lesser extent for Aleppo pine. In conclusion, in the field, holm oak resprouts showed higher competitive ability for P uptake compared to Aleppo pine saplings, but in potted seedlings in common garden conditions this trend was not observed. Therefore holm oak is not always competitively superior to Aleppo pine for P. Potted seedlings of both species had a notable plasticity in shoot/root biomass allocation, but only holm oak increased its proportional allocation to roots when neighbours were present. P availability can be a key factor in growth and competitive relations of these two species, but effects differ depending on soil type, individual age, regeneration type (i.e., seedling versus resprouts), and competitive situation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of far-red light on the growth,mycorrhizas and mineral nutrition of Scots pine seedlings

The effects of supplementary far-red sidelight on the formation of mycorrhizas and on the accumulation and allocation of dry weight and mineral nutrients were studied in Scots pine (Pinus sylvestris L.) seedlings. Starting one week after germination the seedlings were subjected to two different light quality regimes: control and simulated sparse-canopy conditions (FR+). In the FR+ regime, light reflected by neighbouring plants was simulated by means of supplementary far-red light sources, which reduced the horizontal red/far-red photon ratio (R:FR) without affecting PAR. Seedlings were harvested after three months of treatment. FR+ increased stem height and decreased the total dry weight of seedlings. Dry weight allocation to needles was not affected, whereas dry weight allocation to roots was reduced and that to stems was increased in FR+ treated seedlings. The total number of short root tips and developing mycorrhizas per seedling were lower in FR+ than in control plants. Most short roots were developing mycorrhizas, while non-mycorrhizal short roots and mycorrhizas with mantle or external mycelium were very scarce. Changes in the allocation of nutrients in general followed the changes in dry weight allocation, and changes in nutrient content followed those in total dry weight. However, mismatches among these changes resulted in significant changes in nutrient concentrations in some organs: the concentrations of nitrogen and potassium in needles and the concentration of nitrogen in stems were higher in FR+ than in control seedlings. Changes in biomass and nutrient allocation under low R:FR may promote rapid height growth during early development in stands of Scots pine seedlings, but concomitant reductions in growth of the root system and mycorrhizas may negatively affect tree performance over the long term.     

Seedling growth response to added nutrients depends on seed size in three woody genera

1 We tested whether seedlings of small‐seeded species were more reliant on soil nutrients than large‐seeded ones by growing 21 species from three woody genera ( Eucalyptus, Hakea and Banksia ) along a gradient of nutrient availability.
2 At very low nutrient availability, larger seeds produced larger seedlings. This was seen especially among the eucalypts, but the difference was eliminated at optimal soil nutrient levels. Hakea species with large seed mass, and all Banksia species, appeared unable to exploit additional soil nutrients for growth, whatever the level supplied.
3 Larger seeds tended to have proportionately higher contents of N, P and K and, under nutrient‐poor conditions, supplied more of these to their seedlings, although at a diminishing rate.
4 We suggest that large‐seededness could be an adaptation to the high‐light, nutrient‐impoverished habitats in which these species occur by providing the seedling with the mineral nutrients, rather than carbon‐based metabolites, needed for maximizing initial root growth. Reaching reliable moisture before summer (drought avoidance) is an alternative strategy to physiological tolerance of drought.     

磷素营养对降香黄檀幼苗生长及叶片养分状况的影响

设置0、15、30、60、100、150 mg P·株^-1等6个磷素处理开展降香黄檀幼苗盆栽试验,测定各处理幼苗的生长、生物量、叶片养分含量等指标,采用临界浓度法确定降香黄檀幼苗的适宜施磷量,从而探讨不同磷素水平对降香黄檀幼苗生长和叶片养分状况的影响,揭示其磷素需求规律以及适宜的磷供应范围。结果显示磷肥能促进幼苗生长和生物量的积累,而且随施磷量的增加,各指标呈现先升高后降低的趋势,最高值出现在60 mg P·株^-1处理,其苗高、地径、叶面积、生物量分别为对照的3.07、2.35、49.21和24.25倍。施磷显著降低幼苗叶片氮、钾含量,提高磷、镁含量,其中30、60、100 mg P·株^-13个处理间叶片磷含量差异不显著,约为对照的1.65倍。根据幼苗生物量与叶片磷含量、氮钾含量比、磷钾含量比的抛物线关系,确定叶片最适磷含量范围为1.35~2.32 g·kg^-1,由此推断降香黄檀幼苗最适宜的施磷量为60~100 mg P·株^-1。     

Accumulation and Retranslocation of Mineral Nutrients in Developing Needles in Relation to Seasonal Growth of Young Radiata Pine Trees

The growth, accumulation and movement of mineral nutrients (nitrogen,phosphorus, potassium (calcium) and chlorophyll in needles ofyoung radiata pine trees (Pinus radiata D. Don) were examined,from bud break in spring through the following year. Retranslocationof nutrients from needles was measured and is discussed in relationto nutrient requirements for seasonal growth. During the first 4–5 months after bud break when mostneedle growth occurred, all nutrients and chlorophyll accumulatedprogressively, although the concentrations of nitrogen, phosphorusand potassium decreased. During summer, substantial amounts of phosphorus were withdrawnfrom needles less than 6 months old, regardless of positionon the tree and silvicultural practice. In young needles andunder certain environmental conditions, this led to a markedtemporary decline in concentrations, even in fertilized treeson a fertile site. However, the phosphorus content of needleswas quickly restored following autumn rains. Similar fluctuations,including nutrient withdrawal in summer, occurred for nitrogenand potassium, but these were smaller than those observed forphosphorus. Phosphorus was also withdrawn from relatively olderneedles during summer. It was estimated that on a tree basis 86, 48 and 39 per centof the phosphorus, nitrogen and potassium, respectively, insummer shoots could have come from the retranslocation of nutrientsfrom young needles formed during the preceding spring. These results highlight the importance of nutrients stored inneedles to meet the nutrient requirements for growth when environmentalfactors may not be conducive to nutrient uptake from the soil. Pinus radiata D. Don, mineral nutrition, retranslocation, phosphorus, nitrogen, seasonal effects, pine needle growth     

Nutritional and root development factors affecting growth of tissue culture plantlets of loblolly pine

Tissue culture plantlets of loblolly pine ( Pinus taeda L.) were compared to seedlings to quantify growth and developmental differences. The two plant types were grown in containers in a greenhouse and sampled periodically for twenty weeks. Dry weights and nitrogen and phosphorus concentrations of the shoots and roots were determined every two weeks.
During the twenty weeks in the greenhouse, seedlings grew to a greater size than the plantlets, but the relative rates of growth were approximately equal. Plantlets had significantly lower concentrations of nitrogen and phosphorus per g of shoot dry weight. Seedlings were much more efficient at nutrient uptake per g of dry weight of root. Plantlets had thick, unbranched roots, which were inefficient at nitrogen and phosphorus uptake. Nutrient uptake based on an index of root surface area was equal in the plantlets and seedlings.
The main differences between plantlets and seedlings apparently were related to root system morphology rather than physiological processes. The uptake of nutrients showed the greatest difference between the plant types.     

Seed size and seedling growth: differential response of Australian and British Fabaceae to nutrient limitation

Seed size is widely held to exert an important influence over plant establishment, but while large seeds are often assumed to be at an advantage in nutrient-limited conditions, there is in fact, little consistent evidence to support this hypothesis. Here, we examined the interspecific relationship between seedling growth and seed size for Australian and British Fabaceae species in nutrient solutions deficient in nitrogen, phosphorus, potassium or all nutrients combined (distilled water). The British species showed no consistent link between mean seed mass and seedling growth in nutrient-limited conditions. By contrast, all four nutrient-deficient treatments yielded a significant relationship for the Australian species. Linear regression showed that growth under balanced nutrient conditions was positively associated with growth without nutrients, although in fewer cases for the British species. We suggest that habitat-specific differences in regeneration conditions and/or evolutionary history may influence the role that seed size plays in dictating how seedlings of different species respond to nutrient shortage. We recommend caution in attempts to link traits like seed size to wider patterns of plant community ecology.     

The Relationship between Transport of Individual Elements and Dry Matter from the Cotyledons of Pisum sativum L.

The transport of dry matter, nitrogen, potassium, phosphorus,magnesium and sodium from the cotyledons of pea seedlings wasmeasured over a growth period of 4 weeks. A linear relationshipbetween the export of each element and dry matter was observedthroughout the experiment, and it was shown that when a certainproportion of the dry matter available for transport had beenexported the same proportion of the amount of each of the elementsavailable for transport had also moved out of the cotyledons.It is concluded that the movement of an individual solute fromthe cotyledons is controlled by a solute potential gradientexisting between the source and sink.     

Sources of nutrients to rooted submerged macrophytes growing in a nutrient-rich stream

1. The relative contribution of roots and leaves to nutrient uptake by submerged stream macrophytes was tested in experiments where plants were grown in an outdoor flow-channel system. Water was supplied from a nutrient-rich stream with inorganic nitrogen and phosphorus concentrations typical of Danish streams.
2. Four submerged macrophyte species were tested, Elodea canadensis , Callitriche cophocarpa , Ranunculus aquatilis and Potamogeton crispus, and all species were able to satisfy their demand for mineral nutrients by leaf nutrient uptake alone. This was evident from manipulative experiments showing that removal of the roots had no negative impact on the relative growth rate of the plants. Further, the organic N and P concentrations of the plant tissue was constant with time for the de-rooted plants.
3. Enrichment of water and/or sediment had no effect on the relative growth rate of two species, E. canadensis and C. cophocarpa , indicating that in situ nutrient availability was sufficient to cover the needs for growth. Despite the lack of a response in growth rate, a reduced root/shoot biomass ratio was observed with nutrient enrichment of water and/or sediment, and an increased tissue-P concentration in response to open-water enrichment.
4. The open-water nutrient concentrations of the stream in which the experiments were performed are in the upper part of the range found for Danish farmland streams (the majority of Danish streams). Still, however, the negligible effect of nutrient enrichment on the growth of submerged macrophytes observed suggests that mineral nutrient availability might play a minor role in controlling macrophyte growth in most Danish streams.     

施肥对香樟幼苗生长及养分分配的影响

施肥是苗木培育的重要方式,香樟是乡土珍稀阔叶树种,苗木培育对乡土珍稀树种的保护、繁育、推广具有极其重要的作用,施肥对苗木的生长和发育具有重要的影响。因此,为了探讨香樟幼苗生长及植物体内养分分配对施肥的响应,该研究采用正交设计,设置了氮、磷、钾3因素3水平(N、P:0、3、6 g·株-1;K:0、2、4g·株-1),对盆栽香樟幼苗进行指数施肥。结果表明:(1)氮肥对香樟幼苗苗高、地径、生物量的影响最为显著,磷肥和钾肥的影响则较小;(2)氮素在香樟幼苗叶、茎、根中的分布状况主要受氮肥的影响,磷素在香樟幼苗叶、茎、根中的分布状况主要受氮肥和磷肥的影响,钾素在香樟幼苗叶、茎、根中的分布状况主要受钾肥的影响;(3)香樟幼苗的苗高生长与叶片氮含量、叶片磷含量呈显著正相关(P0.05),地径生长与茎氮含量呈显著正相关(P0.05),叶生物量与叶片氮含量、叶片磷含量呈显著正相关(P0.05),茎生物量与叶片磷含量呈显著正相关(P0.05);(4)综合分析得出,对香樟幼苗苗高、地径生长,以及枝叶生物量积累最具促进作用的施肥水平为氮肥(6 g·株-1)、磷肥(6 g·株-1)、钾肥(4 g·株-1)。     

Sediment-related growth limitation of Elodea nuttallii as indicated by a fertilization experiment

1. A fertilization experiment was performed to identify the limiting nutrient for the growth of submerged vegetation in ditches of a peat-grassland system in the Netherlands, in which restoration measures involved ceasing fertilization, exporting nutrients by removal of above-ground plant mass and large-scale introduction of calcium-rich, nutrient-poor artesian water.
2. Growth of Elodea was significantly enhanced by enrichment with nitrogen alone, and by fertilization with nitrogen in combination with phosphorus, and by nitrogen in combination with phosphorus and potassium.
3. Plant tissue nutrient concentrations increased significantly, for nitrogen by enrichment with nitrogen alone, and with nitrogen in combination with phosphorus and potassium; for phosphorus by enrichment with phosphorus alone and with phosphorus in combination with nitrogen and potassium; tissue concentrations of potassium were not enhanced by any treatment.
4. The elemental ratios of treated plants indicated that nitrogen, rather than phosphorus, was limiting in all treatments, except in those involving nitrogen and NK enrichment (when phosphorus was limiting).
5. The efficiency with which plants used nutrients declined with increased supply of nitrogen and phosphorus, but was unchanged when potassium was increased. Efficiencies were similar to those of other aquatic macrophytes.     

Sediment-related growth limitation of Elodea nuttallii as indicated by a fertilization experiment

1. A fertilization experiment was performed to identify the limiting nutrient for the growth of submerged vegetation in ditches of a peat-grassland system in the Netherlands, in which restoration measures involved ceasing fertilization, exporting nutrients by removal of above-ground plant mass and large-scale introduction of calcium-rich, nutrient-poor artesian water.
2. Growth of Elodea was significantly enhanced by enrichment with nitrogen alone, and by fertilization with nitrogen in combination with phosphorus, and by nitrogen in combination with phosphorus and potassium.
3. Plant tissue nutrient concentrations increased significantly, for nitrogen by enrichment with nitrogen alone, and with nitrogen in combination with phosphorus and potassium; for phosphorus by enrichment with phosphorus alone and with phosphorus in combination with nitrogen and potassium; tissue concentrations of potassium were not enhanced by any treatment.
4. The elemental ratios of treated plants indicated that nitrogen, rather than phosphorus, was limiting in all treatments, except in those involving nitrogen and NK enrichment (when phosphorus was limiting).
5. The efficiency with which plants used nutrients declined with increased supply of nitrogen and phosphorus, but was unchanged when potassium was increased. Efficiencies were similar to those of other aquatic macrophytes.     

Effect of flooding with sewage water on three wetland sedges

Plants of three wetland sedges, Carex vesicaria, C. rostrata, and C. gracilis, were subjected to flooding with diluted pig farm sewage water in a sand-culture experiment lasting for one growing season (20 weeks). Sewage water application altered growth dynamics and accumulation of dry matter in all three species; it shifted the distribution of dry matter in favor of above-ground parts; it depressed root growth both in dry mass and length; and it increased percentages of major nutrients (nitrogen, phosphorus, and potassium) in plant dry matter. In the root structure, sewage water application inhibited formation of gas spaces in the cortex, differentiation of exodermis and endodermis, and formation of starch grains in the cortex. The changes in root growth and structure are ascribed to the combination of (1) high nutrient availability, and (2) oxygen deficiency, which developed in the root medium following addition of sewage water. In view of these findings, Carex species are not well suited for use in the treatment of wastewaters rich in both mineral nutrients and organic matter.     

The Circulation of Minerals in Plantations of Pinus sylvestris L.

The dry weights and amounts of sodium, potassium, calcium, magnesium,phosphorus, and nitrogen in the ground flora and organic layersformed over the mineral soil are recorded for an age seriesof plantations of Pinus sylvestris. From these data, the distributionof mineral nutrients within the plantations is deter-mined andcalculations are made of the annual circulation of nutrientsthrough the ecosystem. After 55 years of afforestation the amountof organic matter accumulated over the forest floor is onlyequivalent to the litter fall of the last 6 years. Thus, althoughafforestation results in a progressive accumulation of organicmatter, the woodland community is characterized by a large andrapid transfer of nutrients between the plants and mineral soil.     

Movement of Nutrients in Radiata Pine Needles in Relation to the Growth of Shoots

The pattern of needle growth and the movement of mineral nutrients(nitrogen, phosphorus, potassium and calcium) in needles ofradiata pine (Pinus radiata D. Don) were studied from needleinitiation to 2 years of age. During this period, very prominentcyclic patterns of nutrient accumulation, retranslocation andreplenishment were observed for nitrogen, phosphorus and potassium,which highlighted the potential role of needles as a nutrientreserve for growth. Significant retranslocation occurred from very young needlesabout 3 months after initiation. The phases of retranslocationcoincided with new flushes of shoot growth, and the growth ofnew shoots on a branch resulted in nutrient withdrawal frompre-existing needles, regardless of needle age and season. Suchwithdrawal occurred even in fertilized and irrigated trees onhigh quality sites and under environmental conditions conducivefor nutrient uptake. At all times, except for the short periodafter initiation when needles were actively growing, the nutrientsin the needle were readily available for retranslocation. Contraryto the general view, retranslocation of nutrients was not necessarilyrelated to senescence and ageing of needles. Because new shoots are the primary ‘sinks’ for retranslocatednutrients, an ongoing competition between different parts ofa branch for internal nutrients can be envisaged, preferencebeing for the youngest shoot in the hierarchy. The relevanceof these results to our understanding of ‘optimum nutrition’of pine trees is discussed. Pinus radiata D. Don, radiata pine, mineral nutrition, retranslocation, phosphorus, nitrogen, shoot growth     

Nutrient availability and the carnivorous habit in Utricularia vulgaris

1. Carnivory in plants is thought to enhance growth through an increased supply of nutrients, although there are considerable costs involved. It has been assumed that the relative investment of biomass in traps is inversely proportional to the availability of nutrients from non-carnivorous sources. Our aim was to test the effect of increasing nutrient concentration on investment in carnivory by Utricularia vulgaris .
2. Plants were grown under controlled conditions and nitrogen and phosphorus added at three loadings in a crossed design. Investment in carnivory was assessed as the proportion of (i) leaf biomass and (ii) leaf area comprising traps.
3. There was no effect of nutrient additions on plant growth or periphyton abundance. Investment in carnivory declined with increasing phosphorus loading. There was no effect of nitrogen, despite this being the nutrient commonly thought to be sought by carnivorous plants. Analysis of previously published data also indicated a decline in investment with increasing P availability.
4. Investment in carnivory in U. vulgaris is inversely proportional to the availability of phosphorus from non-carnivorous sources.     

Natural regeneration of selected timber species in the Republic of Congo

Natural regeneration of timber species is critical to the sustainable management of tropical forests. To understand what determines regeneration success of timber species in the Congo Basin, we evaluated whether seedling recruitment rates differed between forest logged 30 years previously and unlogged forest and determined the environmental factors that influence seedling density, growth and survival. We monitored the fate of 2186 seedlings of seven timber species within 462, 25‐m² plots located along 21 transects. We characterized seedling plots by light availability, soil nutrient availability and pH, and abundance of mammalian herbivores and then used linear and generalized linear mixed models to evaluate the variables that influenced seedling density, growth and survival. Light availability and canopy openness were 18% and 81% higher in logged than unlogged forest, and concentration of soil nutrients varied between sites. Seedling density was 32% higher in unlogged than logged forest. Taking all species together, seedling survival was positively correlated with calcium and negatively with magnesium and available phosphorus. Rates of seedling growth increased with available light. Taken separately, seedlings of the selected timber species responded differently to abiotic and biotic factors, demonstrating species‐specific regeneration requirements.     

Studies in the ecology of coastal heath in New South Wales

Effects of heating dry seed were investigated in Banksia serratifolia and B. ericifolia. Seeds were inviable after 7 min exposure at 150°C or higher temperatures but remained viable after exposure at 100°C and lower temperatures. Imbibed seeds show no dormancy, and seeds survive fire insulated from heat in the thick woody follicles which may persist unopened for several years on the parent plant but open following fire. Effects of nutritional supplements were observed in the seed germination and seedling survival and growth of Banksia serratifolia, B. aspleniifolia and B. ericifolia. Germination with radicle and hypocotyls extension was not significantly affected by a wide range of mineral nutrient regimes. In pot experiments using a heath soil seedling survival and growth were adversely affected by moderate additions of phosphate with surviving seedlings showing toxicity symptoms and poor development of proteoid roots. Moderate supplements of potassium had no adverse effect, and in B. serratifolia and B. ericifolia lessened the adverse effects of phosphate addition. The slow growth of the seedlings accords with the normal low nutrient availability in heath soils but seedlings show little capacity for rapid exploitation of any temporary increase of available nutrients after fire.     

Nutrient Supply, Dry-matter Production and Nutrient Uptake of Forest Tree Seedlings

The dry-matter production and nutrient uptake of forest-treeseedlings grown under controlled levels of nutrition in sub-irrigatedsand culture were investigated. Hardwood seedlings attainedgreater dry weight than conifer seedlings, and Pinus radiataa greater dry weight than P. contorta and P. nigra, but thesegreater dry weights were not related to greater leaf/weightratios, although seed size may have been an important influenceon final seedling dry weight. P. radiata was found to have alower requirement for P than P. contorta and P. nigra in oneexperiment. The demand (total nutrient uptake per unit of time) made bythree species for nutrients upon a site was shown to dependmore upon growth-rate per annum than on efficiency (quantityof nutrients required to produce a unit of dry matter). It isconcluded that the ability of tree species to grow satisfactorilyon soils of low nutrient status depends largely on their abilityfor relatively slow growth. On the other hand, species showinggood response to favourable sites may also make satisfactorygrowth on poor sites.     

Tree seedling dynamics in two abandoned tropical dry forests of differing successional status in Yucatán,Mexico: a field experiment with N and P fertilization

The effects of nitrogen (N) and phosphorus (P) fertilization on seedling survival, recruitment, species richness and diversity in two abandoned tropical dry forests (10-yr old, young forest, and c. 60-yr old, old forest) in Yucatán, Mexico, were studied over two years. The seedling dynamics in the control plots were found to be highly seasonal with highest recruitment and lowest death rates during the rainy season. A low percentage of seedlings were resprouts; this important mechanism for forest regeneration had a higher-than-expected survival when compared to seedlings regenerated from seeds. Nutrient addition had significant effects on seedling dynamics in both of the forest regeneration stages. In the young forest, N fertilization facilitated the increase of seedling density. In the old forest, the addition of P decreased seedling diversity, while it increased the recruitment of only a few species. In both forests, P fertilization increased the survival time of seedlings when interacting with light availability and bulk density on the topsoil layer. Results suggest that low nutrient availability combined with low light availability constrain forest succession in Yucatán, Mexico. This revised version was published online in August 2006 with corrections to the Cover Date.