Role of evergreen foliage in the nitrogen economy during shoot growth ofTernstroemia gymnanthera,a warm-temperate broadleaf tree

Accumulation and redistribution of nitrogen were examined during the shoot growth ofTernstroemia gymnanthera, a warm-temperate evergreen broadleaf tree species. Measurements and analyses were confined to the shoot units comprising 2-year-old, 1-year-old and developing current shoots with the foliage of respective ages. Budbreak occurred in early May and nitrogen was rapidly translocated into curent shoots with the progress of their growth. In all of the old organs of the shoot unit, nitrogen concentrations decreased gradually from the time of budbreak to early July. During this period, those old organs supplied more than 60% of the amount of nitrogen needed for the developing current shoots within the same shoot unit. The rest was supplied from the basal organs outside the shoot units comprising branches older than 2 years, stem and roots, by redistribution and/or by absorption from soil. Old leaves, mainly 1-year-old ones, provided about 72% of the total nitrogen derived from the old organs in the shoot units. It was concluded that the evergreen broadleaves served as a large source of nitrogen for the early shoot growth.     

Mycorrhizal infection,phosphorus uptake,and phenology in Ranunculus adoneus: implications for the functioning of mycorrhizae in alpine systems

Phosphorus levels, phenology of roots and shoots, and development of vesicular arbuscular mycorrhizal (VAM) fungi were monitored for two years in natural populations of the perennial alpine herb, Ranunculus adoneus. The purpose of this study was to understand how phosphorus uptake relates to the phenology of R. adoneus and to ascertain whether arbusculus, fungal structures used for nutrient transfer, were present when maximum phosphorus accumulation was occurring. Arbuscules were only present for a few weeks during the growing season of R. adoneus and their presence corresponded with increased phosphorus accumulation in both the roots and shoots of R. adoneus. In addition, phosphorus accumulation and peaks in mycorrhizal development occurred well after plant reproduction and most plant growth had occurred. The late season accumulation of phosphorus by mycorrhizal roots of R. adoneus is stored for use during early season growth and flowering the following spring. In this way R. adoneus can flower before soils thaw and root or mycorrhizal nutrient uptake can occur.     

Biology of the bamboo Chusquea culeou (Poaceae: Bambusoideae) in southern Argentina

Over a period of 7 years the biology and phenotypic variability of Chusquea culeou were studied at 5 locations in cool temperate forests of southern Argentina. Excavated rhizomes had an average of 1.1 successful rhizome buds, and an average of 2.1 years elapsed between successive generations of rhizomes. Rhizome buds usually develop within the first four years after a rhizome forms. Height, volume and weight of a culm can be calculated from its diameter 1 m above the ground. Culm size, length of foliage leaf blades, and pattern of secondary branching differed among study sites. Dead culms were numerous and commonly remained erect for more than 7 years after dying. New culm shoots appear in spring and reach full size within a few months. Shoots can grow more than 9 cm/day. Less than half of the shoots survived a year; most were killed by moth larvae. Multiple primary branch buds emerge through the culm leaf sheaths in the second spring. The mean number of branch buds at mid-culm nodes varied between 34.8 and 81.5, and the mean number of primary branches was between 22.8 and 40.8. Number and length of branches, and number and length of foliage leaf blades at each node is related to the position of the node on a culm. Most branches grow about 3 cm and produce 1 to 3 foliage leaves annually. Foliage leaf blades generally live 2 years or more; few survive 6 years. Relative lengths of foliage leaf blades and their spacing along a branch permit recognition of annual cohorts.Both gregarious and sporadic flowering have been reported, and every year a few isolated plants flower and die. Length of the life cycle is unknown. Seedlings require up to 15 years to produce culms of mature size. Foliage branches may live more than 23 years, and culms may survive 33 years. Extensive loss of new shoots to predation suggests that gregarious flowering may be driven by a need to escape parasitism. C. culeou clumps expand slowly. Average annual rate of increase of the number of live culms in a clump was 4.6%. Methods of seed dispersal are undocumented. A dense stand of Chusquea culeou had an estimated phytomass of 179 tons/hectare (dry weight), 28% of which was underground. Net annual production was about 16 t/ha dry weight.     

Variation in foliar nutrients in Eucalyptus trees in eastern and Western Australia

Abstract Levels of nitrogen, phosphorus and potassium were measured for the foliage of two co-dominant eucalypts at each of two sites, one in eastern Australia and the other in Western Australia. In eastern Australia, foliage was sampled in the canopy and subcanopy for narrow-leaved ironbark Eucalyptus crebra and grey box E. mollucana and in Western Australia, for jarrah E. marginata and marri E. calophylla. The Western Australian trees were also sampled for ‘young’ and ‘old’ leaves. Both eucalypts in eastern Australia had greater nitrogen and phosphorus levels, but lower potassium, than E. marginata or E. calophylla. Eucalyptus calophylla foliage had greater levels of all three nutrients than E. marginata foliage as did E. crebra relative to E. mollucana. At both sites, foliar nutrient levels were greater in the canopy than subcanopy foliage, and, at least in Western Australia, the younger leaves had greater nutrient levels than the older leaves. The observed differences in foliar nutrient levels are consistent with observed trends in the abundance and diversity of foliage arthropods and the use of the trees as foraging substrates by birds.     

The effect of previous defoliation of pole-stage lodgepole pine on plant chemistry,and on the growth and survival of pine beauty moth (Panolis flammea) larvae

Summary A study of the effects of defoliation by insects on the chemistry of lodgepole pine (Pinus contorta), and on the performance of Panolis flammea (Lepidoptera; Noctuidae) larvae, was carried out in a forest in northwest Scotland I year after a severe outbreak of P. flammea had caused extensive defoliation. Larval weight and survival were not significantly different on trees that had experienced different levels of defoliation in 1986. The nitrogen and tannin content of current and previous years' pine needles was not significantly affected by defoliation (although both were slightly greater in the foliage of defoliated trees). Phosphorus content of young pine foliage was lower (but not significantly lower except on one occasion) on heavily defoliated trees. On all sampling occasions, however, the nitrogen: phosphorus ratio was significantly higher on heavily defoliated trees. There were large differences in monoterpene composition of the previous year's shoots associated with defoliation intensity, but these differences had largely disappeared in the new growth. The results are discussed in relation to other studies on the effects of insect damage on plant chemistry and insect performance and in relation to the abundance of P. flammea in Scotland.     

The chemical composition of pine foliage in relation to the population dynamics of the pine beauty moth,Panolis flammea,in Scotland

Summary This paper reports part of a study to determine why damaging outbreaks of the pine beauty moth, Panolis flammea (D & S) (Lepidoptera: Noctuidae), in Scotland are frequent on lodgepole pine but do not occur on Scots pine, and why outbreaks on lodgepole pine are mainly confined to trees growing in deep unflushed peat. The elongation of shoots and the growth of needles of Scots pine occurred later in the season than did those of lodgepole pine. The foliage of Scots pine generally had a higher level of nitrogen, and consistently had a higher level of phosporus, but had a consistently lower level of tannins than that of lodgepole pine during the period when the larvae were feeding each year. The nitrogen content of the foliage of lodgepole pine growing in an iron pan soil was generally higher than that of lodgepole pine growing in deep peat during the same period but there were no general differences in the phosphorus or tannin contents of lodgepole pine in the 2 soil types. These findings suggest that Scots pine is a more suitable host plant than lodgepole pine and that the foliage of lodgepole pine growing in deep peat is not more suitable than lodgepole pine growing in an iron pan soil. On the basis of the chemical analyses used in this study, it is concluded that the abundance of pine beauty moth in Scotland is not strongly influenced by the nutritional suitability of its host plants.     

Seasonal patterns of nutrient deposition in a Quercus douglasii woodland in central California

The monthly deposition of total nitrogen, phosphorus, potassium, calcium and magnesium via canopy throughfall, and various components of the litterfall was measured for 31 months under mature Quercus douglasii and in the bulk precipitation in the surrounding open grassland. Seasonal patterns of nutrient concentration in leaf litter, throughfall, and precipitation were also measured. Total annual subcanopy deposition exceeded open precipitation deposition by approximately 45–60x for nitrogen, 5–15x for phosphorus, 30–35x for potassium, 25–35x for calcium, and 5–10x for magnesium. Total annual subcanopy deposition was low in comparison to other oak woodland sites reported in the literature. Throughfall and leaf litter were the primary sources of nutrients and thus determined the seasonal peaks of nutrient deposition. The first autumn rains and leaf fall were associated with one peak in nutrient deposition, and throughfall during early spring leaf emergence was associated with a second peak in potassium, magnesium and phosphorus. Non-leaf plant litter (excluding acorns) provided approximately 15–35% of most nutrients, with twigs and bark depositing over 12% of the annual calcium flux in 1987–1988, and flower litter depositing over 8% of the annual nitrogen flux in 1986–1987. Acorns had high concentrations of phosphorus and nitrogen and during the mast season of 1987–1988 they contained a large proportion of the total subcanopy annual flux of these elements. With acorns excluded, total annual nutrient deposition was similar between years, but timing of nutrient deposition differed. Late summer leaf fall associated with drought, variation in precipitation, and variation in deposition of non-leaf parts were associated with seasonal differences in nutrient deposition between years.     

Growth ofAgrostis capillaris andDeschampsia flexuosa in relation to decomposition of and nutrient release by Sitka spruce litter

Summary Bags containing Sitka spruce litter (0, 15, 50 g) were placed in flower pots and covered with sand. Pots were watered at weekly intervals with nutrient solutions with and without nitrogen and with and without phosphorus. Decomposition was measured by carbon dioxide evolution from pots without plants. Neither added nitrogen nor added phosphorus had any marked effect on the rate of decomposition, which amounted to 14% loss of carbon in a year. The two grass species responded similarly to fertilizer; 72% of added nitrogen and 90% of added phosphorus were recovered in plant parts.A. capillaris captured nutrients more effectively from spruce litter than didD. flexuosa, recovering 13% of the phosphorus in the litter but only 5% of the nitrogen. Neither uptake of nitrogen nor uptake of phosphorus was enhanced in plants receiving fertilizer additions of the other nutrient.     

Interplant variation in creosotebush foliage characteristics and canopy arthropods

Summary We conducted a field study to test the hypothesis that creosotebush (Larrea tridentata) shrubs growing in naturally nutrient-rich sites had better quality foliage and supported greater populations of foliage arthropods than shrubs growing in nutrient-poor sites. Nutrient-rich sites had significantly higher concentrations of soil nitrogen than nutrient-poor sites. Multivariate analysis of variance revealed significant differences between high nutrient and low nutrient shrubs based on a number of structural and chemical characteristics measured. High nutrient shrubs were larger, had denser foliage, greater foliage production, higher concentrations of foliar nitrogen and water, and lower concentrations of foliar resin than low nutrient shurbs. Numbers of foliage arthropods, particularly herbivores and predators, were significantly higher on high nutrient shrubs. Shrub characteristics and foliage arthropod abundances varied considerably from shrub to shrub. Shrub characteristics representing shrub size, foliage density, foliage growth, and foliar nitrogen and water concentrations were positively correlated with arthropod abundances. Foliar resin concentrations were negatively correlated with foliage arthropod abundances. The positive relationship between creosotebush productivity and foliage arthropods is contradictory to the tenet that physiologically stressed plants provide better quality foliage to insect herbivores.     

Seasonal changes in element contents in mangrove element retranslocation during leaf senescene

The concentrations of nitrogen (N), phosphorus (P), potassium (K), sodium (Na), calcium (Ca), magnesium (Mg) and chlorine (Cl) were followed monthly in pre-senescence leaves and post-abscission leaves of Kandelia candel (L.) Druce at the Jiulongjiang estuary, and Fujian, China. The element retranslocation efficiency (RE) was studied during leaf senescence. The element RE's evaluated using different methods were compared and a new method was put forward to evaluate element RE during leaf senescence in evergreen trees without concentrated leaf fall. The results showed that during leaf senescence, 77.22% N, 57.53% P, and 44.51% K were translocated out of senescing leaves. Translocation of nutrients out of senescing leaves back into shoots was an important nutnent-conservation mechanism for N and P, was less important for K, and did not occur for Ca, Mg, Na, or Cl. One of the reasons for the high primary productivity of mangroves in nutrient poor sites (especially with low N) is the high nutrient use efficiency.     

Plant biomass and nutrient levels of a tropical macrophyte (Cyperus papyrus L.) receiving domestic wastewater

The aboveground biomass and nutrient content ofCyperus papyrus were determined in a small tropical swamp receiving domestic wastewater. It was found that the biomass (4,955 g.m^–2 dry weight) was the highest ever reported for papyrus. The levels of both nitrogen and phosphorus in the plant organs were very high. The nitrogen concentrations of the various plant organs were 4.8% roots, 8.4% rhizomes, 4.5% scales, 4.8% culms, and 6.2% umbels on dry weight basis. As to phosphorus the concentrations were 0.09% roots, 0.11% rhizomes, 0.09% scales, 0.10% culms, and 0.13% umbels. The high biomass and nutrient contents of the plants may have been caused by the high nutrient levels in the surrounding water. Comparison of the nitrogen to phosphorus ratios in the plants to those in the surrounding water showed that the plants stored very high amounts of nitrogen.     

The impact of defoliation on the foliar chemistry of southern rātā (

Brushtail possums (Trichosurus vulpecula) tend to eat young canopy foliage in southern rätä (Metrosideros umbellata), and browsing tends to be concentrated on only a few trees. Samples collected as part of an artificial defoliation experiment were analysed for NPK (nitrogen, phosphorus, and potassium), carbohydrate, and polyphenolic concentrations to determine whether changes in foliar chemistry associated with defoliation provide an explanation for these patterns of browsing. In non-defoliated trees, NPK concentrations were highest in young leaves and declined with age, while concentrations of carbohydrates and polyphenolics were independent of leaf age. Nitrogen, phosphorus and polyphenolic concentrations were consistently higher in canopy (sun) versus subcanopy (shade) foliage regardless of leaf age, a trend that was reversed for potassium. Partial (50%) defoliation had little effect on foliar chemistry, regardless of its timing. Total (100%) defoliation stimulated NPK concentrations and depressed condensed tannin concentrations of new foliage produced by the surviving shoots. These results suggest that brushtail possums may focus their feeding on only a few trees because of nutritional changes to leaves as a result of browsing.     

Carbon,phosphorus and nitrogen budgets of the littoral Equisetum belt in an oligotrophic lake

Stores and flows of carbon, phosphorus and nitrogen in a littoral Equisetum stand were studied in 1978–1980 in the oligotrophic, mesohumic lake Pääjärvi, southern Finland. The major carbon and nutrient stores were sediment and Equisetum. The seasonal cycle of the macrophyte vegetation had a profound influence on the whole littoral ecosystem. In spring, when only dead remains of Equisetum were present above ground, there were few differences in nutrient, chlorophyll a and zooplankton concentrations between the littoral and the open lake; phytoplankton and epiphytes were the major producers.In early June, when new shoots of Equisetum reached the water surface, water exchange between the littoral and the open lake started to diminish, and the characteristic features of a closed macrophyte zone gradually developed: by August the P, Chl a and zooplankton concentrations in the littoral were 5–10 times those in the open lake. From late June until autumn Equisetum was overwhelmingly dominant both in biomass and in production.The measured total primary production and respiration values indicated a high rate of internal cycling of carbon and nutrients. The daily P requirements of plant growth exceeded the total P stored in the water by a factor of 2–4, and also exceeded the release of nutrients in excretion. High N:P ratios in the water (total 10–64, inorganic 18–171) suggested that P was probably always the limiting nutrient.The P content of the annual production of Equisetum in Pääjärvi was 2.3% of the mean annual P load, and 5.3% of the mean total P storage in the water volume of the lake.     

The resource balance of Milium effusum with emphasis on environmental resource supply

Summary Growth of the broad-leaved graminoid Milium effusum, occurring in shady deciduous forests, was matched with periods of high light influx through the tree canopy in spring and autumn. Fertile shoots grew faster than sterile shoots. Leaves on flowering shoots were fully developed when the budbreak started on the trees, whereas nonflowering shoots had fully developed leaves when the tree canopy closed. Leaf concentrations of N and P were high (6.1 and 0.74% respectively) in spring but decreased as the leaves expanded. Maximum pool sizes of N and P in whole tillers were reached about one month after the onset of spring growth, whereas maximum spring pools of K, Mg, and Ca were timed with peak biomass about one month later. The leaves lost nutrients during summer when no growth took place. Since leaching losses were negligible, nutrients were probably allocated from the leaves to support root growth. Autumn reallocation to winter stores was low. The pattern of growth and nutrient use suggests that light availability, i.e., the resource in relatively lowest supply, regulates the investment of the resource in highest supply, i.e., nutrients. This is consistent with previously reported observations on Eriophorum vaginatum, a graminoid of low nutrient — high light environments. This species utilizes nutrients efficiently at the expense of less efficient acquisition of carbon. We suggest that selection for efficient utilization of the resource in lowest relative supply has been a strong driving force behind the physiological adaptation of both species to their environments.     

Effects of self-thinning of shoots on the nutrient budgets of Zizania latifolia

This experimental study focused on the seasonal changes and mobilization of nutrients between plant parts to understand the implications of self-thinning in the ecology of Zizania latifolia (Griseb.) Turcz. ex Stapf. The observations of shoot density, above- and belowground biomass, and total nitrogen and total phosphorus concentrations were conducted from February 2002 to August 2003. The biomass of shoots that died during the period and daily uptake of nutrients were determined. The shoot density sharply increased until mid-April and thereafter decreased significantly due to self-thinning of shoots. Total nitrogen and phosphorus concentrations of rhizomes decreased initially due to translocation to new shoots; however, the nutrients of rhizomes were slightly replenished from dead shoots during self-thinning. In contrast to other species, self-thinning of Z. latifolia shoots reallocates some of the minerals contained in the dead shoots back to the rhizomes, which can be regarded as a strategy to replenish the reduced resources of the rhizomes. The initial intensive growth of shoots can be regarded as a strategy to maintain resource competition.     

Phytophagous insects enhance nitrogen flux in a desert creosotebush community

Summary We tested the hypothesis that herbivorous insects on desert shrubs contribute to short-term nitrogen cycling, and increase rates of nitrogen flux from nutrient rich plants. Creosotebush (Larrea tridentata) shrubs were treated with different combinations of fertilizer and water augmentations, resulting in different levels of foliage production and foliar nitrogen contents. Foliage arthropod populations, and nitrogen in canopy dry throughfall, wet throughfall and stemflow were measured to assess nitrogen flux rates relative to arthropod abundances on manipulated and unmanipulated shrubs over a one-month period during peak productivity. Numbers and biomass of foliage arthropods were significantly higher on fertilized shrubs. Sap-sucking phytophagous insects accounted for the greatest numbers of foliage arthropods, but leaf-chewing phytophagous insects represented the greatest biomass of foliage arthropods. Measured amounts of bulk frass (from leaf-chewing insects) were not significantly different among the various treatments. Amounts of nitrogen from dry and wet throughfall and stemflow were significantly greater under fertilized shrubs due to fine frass input from sap-sucking insects. Increased numbers and biomass of phytophagous insects on fertilized shrubs increased canopy to soil nitrogen flux due to increased levels of herbivory and excrement. Nitrogen excreted by foliage arthropods accounted for about 20% of the total one month canopy to soil nitrogen flux, while leaf litter accounted for about 80%.     

Growth and production ofPhragmites australis in Lake Vechten (the Netherlands)

Summary A study was carried out on several production characteristics ofPhragmites australis stands in Lake Vechten, the Netherlands. The aim was to quantify the role of this plant species in the lake's primary production and nutrient cycles (C, N and P).Maximum biomass values of 1247 and 286 g org.wt.m^–2 were measured in, respectively, shoots and underground organs in August. The shoot: root ratio was always>1. Plant litter accumulated particularly in spring and autumn. The relative growth rates of early shoots differed from those determined for the whole vegetation, indicating shoots of different ages in the latter. This was confirmed by counting the number of plants.The chemical composition of the plants changed with season. A carbon peak as high as 50.94% org.wt. was found in spring, which coincided with a nitrogen peak of 2.88% org. wt. At the end of the season N dissapeared faster from the shoots than C, whereas P declined from spring onwards.The production ofPhragmites contributed in 1979 for 5.8% to the lake's total primary production expressed in C, whereas the share of the total macrophytic unit amounted to 8.7%.The potential role ofPhragmites in the lake's nutrient cycles was largely due to nutrient leakage and decomposition.     

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     

Seasonal dynamics of shoot nutrients in Schoenus ferrugineus (Cyperaceae)

Seasonal patterns of biomass and standing stock (g m⁻²) of nitrogen, phosphorus, potassium and calcium were documented in Schoenus ferrugineus L. from an annually mowed calcareous fen in southern Germany. The peak standing crop of nitrogen, phosphorus and potassium occurred in mid-July. Prior to the end of the growing season, 44% of total potassium, 34% of total phosphorus and 28% of total nitrogen were retranslocated out of the shoots, based on differential nutrient losses between covered and uncovered plots. There is no evidence that any calcium was retranslocated, but 24% of peak calcium and 13% of potassium was leached from standing dead shoots. More than half of the peak phosphorus was found in the overwintering green shootbases. In the light of the low availability of nutrients in the soil, retranslocation of phosphorus in particular is essential, if adequate internal nutrient levels are to be maintained, despite various proposed uptake mechanisms.     

Mineral composition and growth of Colorado spruce (Picea pungens) seedlings under calcareous soil conditions

Summary Field studies were conducted to assess the mineral nutrition and growth of Colorado spruce (Picea pungens Engln) seedlings (2–4 yr) from provenances selected for superior growth on calcareous prairie soils. Tissue nutrient concentrations and response to nitrogen were determined by use of foliar analysis and growth pattern studies. Soil conditions ranged from 7.6–7.8 for pH, 12–23% for total CaCO₃, and 5–6% for active CaCO₃. Foliage mineral composition showed relatively low phosphorus (0.09–0.15%) and high calcium (0.45–1.52%) assimilation. Seasonal growth and seedling response to added nitrogen was not adversely affected by the calcareous soil condition. Levels of nitrogen in the foliage required for optimum growth ranged from 1.5–2.0% and were similar to that of other conifer species.