Interactions between leaf nitrogen status and longevity in relation to N cycling in three contrasting European forest canopies

Seasonal and spatial variations in foliar nitrogen (N) parameters were investigated in three European forests with different tree species, viz. beech (Fagus sylvatica L.), Douglas fir (Pseudotsuga menziesii, Mirb., Franco) and Scots pine (Pinus sylvestris L.) in Denmark, The Netherlands and Finland, respectively. This was done in order to obtain information about functional acclimation, tree internal N conservation and its relevance for both ecosystem internal N cycling and foliar N exchange with the atmosphere. Leaf N pools generally showed much higher seasonal variability in beech trees than in the coniferous canopies. The concentrations of N and chlorophyll in the beech leaves were synchronized with the seasonal course of solar radiation implying close physiological acclimation, which was not observed in the coniferous needles. During phases of intensive N metabolism in the beech leaves, the NH4+ concentration rose considerably. This was compensated for by a strong pH decrease resulting in relatively low Γ values (ratio between tissue NH4+ and H+). The Γ values in the coniferous were even smaller than in beech, indicating low probability of NH3 emissions from the foliage to the atmosphere as an N conserving mechanism. The reduction in foliage N content during senescence was interpreted as N re-translocation from the senescing leaves into the rest of the trees. The N re-translocation efficiency (ηr) ranged from 37 to 70% and decreased with the time necessary for full renewal of the canopy foliage. Comparison with literature data from in total 23 tree species showed a general tendency for ηr to on average be reduced by 8% per year the canopy stays longer, i.e. with each additional year it takes for canopy renewal. The boreal pine site returned the lowest amount of N via foliage litter to the soil, while the temperate Douglas fir stand which had the largest peak canopy N content and the lowestηr returned the highest amount of N to the soil. These results support the hypothesis that a high N status, e.g. as a consequence of chronically high atmospheric N inputs, increases ecosystem internal over tree-bulk-tissue internal N cycling in conifer stands. The two evergreen tree species investigated in the present study behaved very differently in all relevant parameters, i.e. needle longevity, Nc and ηr, showing that generalisations on tree internal vs. ecosystem internal N cycling cannot be made on the basis of the leaf habit alone.

Needle longevity as a criterion of response to a climatic fluctuation (so called heat wave) in Scots pine populations at early phases of ontogeny

The paper presents results of an assessment of needle age classes in Scots pine partial populations in the period of three and five years after outplanting in a lowland site in relation to conditions created by climatic elements. The method is based on Skuhravý’s (1987) description when the state of needle age classes of the third whorl (from the apex) is evaluated just once. Five-year summary of meteorological data was assessed according to the criteria of determination of periods of heat waves and dry spells. Differences in numbers of living needle age classes in the groups of Scots pine populations in the period of three and five years after their outplanting in permanent lowland sites proved the highest total number of living needle classes in the group of local lowland populations. On the contrary, the lowest total number of living needle classes was found out in the group of mountain populations. Critical climatic periods are overlaps of heat waves and dry spells that illustrate the complexity of moisture and temperature conditions of lowland sites. Although the Scots pine is a xerophytic species, it responded to the cumulation of adverse factors by accelerated needle senescence. Its physiological response to heat and dry stress consists in needle shedding, i.e. reduction in the assimilating area. As mountain populations are not usually exposed to heat waves, the impact of this phenomenon on them was the highest. On the other hand, the local (lowland) populations carrying information on the patterns of local climate from the reproduction period do not undergo any marked reduction in needle age classes.  

Variation in Scots pine needle longevity and nutrient conservation in different habitats and latitudes

Within-species variation in needle traits is an important characteristic of conifers enabling trees to grow in different environments. We compared mean needle age (NA), shoot growth, and nutrient conservation in Scots pine ( Pinus sylvestris L.) populations in different habitats and latitudes (peatlands and Vaccinium -type stands in Estonia (59°N) and Lapland (66°30′N)). In Vaccinium-type stands, trees with higher NA (mostly in Lapland) had lower shoot length increment and lower concentration of nitrogen and phosphorus in current-year needles than trees with low NA (mostly in Estonia). However, in peatlands, variation in NA (which was as high as in the Vaccinium-type stands) was weakly or insignificantly related to shoot growth and needle nutrient concentration. Within latitudes, pines with different shoot length increments and needle nutrient concentrations tend to have similar NAs. Resorption efficiency and concentration of nitrogen and phosphorus in senescent needles decreased with the initial concentrations of these nutrients in green needles. Our results demonstrate that slow growth and low needle nutrient concentration are not necessarily followed by higher NA and greater nutrient conservation in Scots pine. This is the opposite of the results often obtained in among-species comparisons or within species along latitudinal and altitudinal gradients.

Effects of repeated fertilization on needle longevity, foliar nutrition, effective leaf area index, and growth characteristics of lodgepole pine in interior British Columbia, Canada

We investigated the effects of repeated fertilization (either periodically every 6 years or annual fertilization) on needle longevity and growth response in two juvenile lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) stands in the interior of British Columbia, Canada. Annual fertilization decreased needle longevity by 23% at the Kenneth Creek site and by 30% at Sheridan Creek, compared with the control treatments at each site. At Sheridan, repeated fertilization significantly increased effective leaf area index, foliated shoot length, and annual shoot growth. However, none of these variables was significantly altered by repeated fertilization at Kenneth. At both locations, fertilization elevated nutrient concentrations in the current year's foliage. Annual fertilization increased nitrogen concentration in mid-crown branches of retained cohorts (1998–2002) at both study sites. Furthermore, annual nitrogen addition apparently induced or exacerbated copper and iron deficiency in these stands, especially at Kenneth Creek, which may be related to the premature loss of foliage. Nutrient imbalance may also be related to poor effective leaf area index and growth response at Kenneth Creek. Stem growth efficiency declined with annual fertilization at Kenneth Creek because of accelerated turnover of needles, increased allocation of growth to branches, and probably reduced photosynthetic capacity.