994 resultados para Scots Pine
Resumo:
Ultrasonic sound velocity measurements with hand-held equipment remain due to their simplicity among the most used methods for non-destructive grading of sawn woods, yet a dedicated normalization effort with respect to strength classes for Spanish species is still required. As part of an ongoing project with the aim of definition of standard testing methods, the effect of the dimensions of commonly tested Scots pine (Pinus sylvestris L.) timbers and equipment testing frequency on ultrasonic velocity were investigated. A dedicated full-wave finite-difference time-domain software allowed simulation of pulse propagation through timbers of representative length and section combinations. Sound velocity measurements vL were performed along the grain with the indirect method at 22 kHz and 45 kHz for grids of measurement points at specific distances. For sample sections larger than the cross-sectional wavelength ?RT, the simulated sound velocity vL converges to vL = (CL/?)0.5. For smaller square sections the sound velocity drops down to vL = (EL/?)0.5, where CL, EL and ? are the stiffness, E-modul and density, respectively. The experiments confirm a linear regression between time of flight and measurement distance even at less than two wavelength menor que2?L distance, the fitted sound speed values increased by 15% between the two tested frequencies.
Resumo:
In mixed stands, inter-specific competition can be lower than intra-specific competition when niche complementarity and/or facilitation between species prevail. These positive interactions can take place at belowground and/or aboveground levels. Belowground competition tends to be size symmetric while the aboveground competition is usually for light and almost always size-asymmetric. Interactions between forest tree species can be explored analyzing growth at tree level by comparing intra and inter-specific competition. At the same time, possible causes of niche complementarity can be inferred relating intra and inter-specific competition with the mode of competition, i.e. size-symmetric or sizeasymmetric. The aim of this paper is to further our understanding of the interactions between species and to detect possible causes of competition reduction in mixed stands of beech (Fagus sylvatica L.) with other species: pine?beech, oak?beech and fir?beech. To test whether species growth is better explained by size-symmetric and/or size-asymmetric competition, five different competition structures where included in basal area growth models fitted using data from the Spanish National Forest Inventory for the Pyrenees. These models considered either size-symmetry only (Reineke?s stand density index, SDI), size-asymmetry only (SDI of large trees or SDI of small trees), or both combined. In order to assess the influence of the admixture, these indices were introduced in two different ways, one of which was to consider that trees of all species compete in a similar way, and the other was to split the stand density indices into intra- and inter-specific competition components. The results showed that in pine?beech mixtures, there is a slightly negative effect of beech on pine basal area growth while beech benefitted from the admixture of Scots pine; this positive effect being greater as the proportion of pine trees in larger size classes increases. In oak?beech mixtures, beech growth was also positively influenced by the presence of oaks that were larger than the beech trees. The growth of oak, however, decreased when the proportion of beech in SDI increased, although the presence of beech in larger size classes promoted oak growth. Finally, in fir?beech mixtures, neither fir nor beech basal area growth were influenced by the presence of the other species. The results indicate that size-asymmetric is stronger than size-symmetric competition in these mixtures, highlighting the importance of light in competition. Positive species interactions in size-asymmetric competition involved a reduction of asymmetry in tree size-growth relationships.
Unravelling past flash flood activity in a forested mountain catchment of the Spanish Central System
Resumo:
Flash floods represent one of the most common natural hazards in mountain catchments, and are frequent in Mediterranean environments. As a result of the widespread lack of reliable data on past events, the understanding of their spatio-temporal occurrence and their climatic triggers remains rather limited. Here, we present a dendrogeomorphic reconstruction of past flash flood activity in the Arroyo de los Puentes stream (Sierra de Guadarrama, Spanish Central System). We analyze a total of 287 increment cores from 178 disturbed Scots pine trees (Pinus sylvestris L.) which yielded indications on 212 growth disturbances related to past flash flood impact. In combination with local archives, meteorological data, annual forest management records and highly-resolved terrestrial data (i.e., LiDAR data and aerial imagery), the dendrogeomorphic time series allowed dating 25 flash floods over the last three centuries, with a major event leaving an intense geomorphic footprint throughout the catchment in 1936. The analysis of meteorological records suggests that the rainfall thresholds of flash floods vary with the seasonality of events. Dated flash floods in the 20th century were primarily related with synoptic troughs owing to the arrival of air masses from north and west on the Iberian Peninsula during negative indices of the North Atlantic Oscillation. The results of this study contribute considerably to a better understanding of hazards related with hydrogeomorphic processes in central Spain in general and in the Sierra de Guadarrama National Park in particular.
Resumo:
Ultrasound wave velocity was measured in 30 pieces of Spanish Scots pine (Pinus sylvestris L.), 90 x 140 mm in cross-section and 4 m long. Five different sensor placement arrangements were used: end to end (V0), face to opposite face, edge to opposite edge, face to same face and edge to same edge. The pieces were successively shortened to 3, 2 and 1 m, in order to obtain these velocities and their ratios to reference value V0 for different lengths and angles with respect to the piece axis for the crossed measurements. The velocity obtained in crossed measurements is lower than V0. A correction coefficient for crossed velocities is proposed, depending on the angle, to adjust them to the V0 benchmark. The velocities measured on a surface, are also lower than V0, and their ratio with respect to V0 is close to 0.97 for distances equal to or greater than 18 times the depth of the beam.
Resumo:
La fase de establecimiento del regenerado es un proceso crítico para el desarrollo posterior de la masa tanto por las elevadas tasas de mortalidad que habitualmente lleva asociadas, como por proporcionar el material de partida del que van a disponer las fases subsiguientes. Las restricciones a la germinación y establecimiento de la regeneración del pino silvestre varían enormemente entre las distintas regiones de su extensa área de distribución geográfica. La región Mediterránea constituye un hábitat marginal de la especie en el que las condiciones ecológicas son muy diferentes a las del grueso de su área de distribución. Frente a otras limitaciones (frío, luz, encharcamiento…), en el entorno mediterráneo la tasa de mortalidad parece estar asociada a las condiciones micrometeorológicas del período estival - particularmente, a la sequía -, así como a la presencia excesiva de ganado o ungulados silvestres. No obstante, la mayoría de la información disponible sobre el proceso de regeneración de la especie procede del centro y norte de Europa, por lo que no es de aplicación directa en nuestra región, en la que los estudios de este tipo son mucho más escasos. El presente trabajo pretende contribuir a paliar esta relativa escasez a través del estudio del proceso de regeneración natural en el monte “Cabeza de Hierro”, masa irregular por bosquetes de pino silvestre, paradigma de gestión sostenible y uso múltiple. En este entorno, se pretende caracterizar y cuantificar tanto el proceso de germinación y supervivencia de la especie como la influencia de la cobertura vegetal (estratos arbóreo, arbustivo y herbáceo, y capa de restos vegetales) en su desarrollo. Se persigue así mismo analizar el efecto de la compactación del suelo sobre la persistencia de la masa y contrastar y comparar la eficacia de dos tratamientos edáficos de ayuda a la regeneración: escarificado y decapado+acaballonado. Con este fin se han planteado dos diseños experimentales consistentes en sendas redes de muestreo (Red de Muestreo I o RM I y Red de Muestreo II o RM II) integradas, respectivamente, por 192 y 24 parcelas de 1,5x1,5 m ubicadas bajo distintas condiciones de cobertura vegetal. Sobre una parte de estas parcelas (1/4 en la Red de Muestreo I; 1/2 en la Red de Muestreo II) se han aplicado tratamientos de ayuda a la regeneración (RM I: escarificado; RM II: decapado+acaballonado) y, tras llevar a cabo siembras controladas al inicio del período vegetativo, se han practicado controles periódicos de germinación y supervivencia durante uno (RM II) y tres años consecutivos (RM I). Se han realizado así mismo mediciones complementarias de variables micrometeorológicas, espesura, recubrimiento superficial del suelo y compactación. Los resultados obtenidos a partir de las experiencias realizadas en el monte objeto de estudio permiten concluir que, en relación con el proceso de regeneración natural de la especie en este tipo masa y entorno: 1) la regeneración del pino silvestre durante el primer período vegetativo presenta una tasa de éxito muy baja (1,4% de los sembrados), provocada por una elevada mortalidad durante el primer período estival (>92%) subsiguiente a una germinación de en torno al 17% de las semillas viables que llegan al suelo; 2) la mortalidad sigue siendo elevada hasta el tercer período vegetativo, en que comienza a reducirse significativamente hasta alcanzar el 45%; 3) la cobertura vegetal influye significativamente tanto en el proceso de germinación como en el de supervivencia, aunque ambos procesos presentan una baja correlación linear que pone de manifiesto que los lugares idóneos para la germinación no siempre son los más adecuados para la supervivencia; 4) la escarificación del suelo mejora las tasas iniciales de germinación y supervivencia, pero empeora la tasa de supervivencia posterior (años 2 y 3), por lo que su efecto a medio plazo no resulta significativo; 5) el decapado+acaballonado presenta mejores resultados que el escarificado durante el primer verano, aunque sólo resulta efectivo en condiciones intermedias de espesura de masa; 6) la compactación edáfica no resulta limitante para la productividad ni la persistencia de la masa considerada. ABSTRACT Seedling establishment is critical for later stand progress because it involves high mortality rates and the surviving saplings constitute the starting material for all the subsequent stages. Restrictions for Scots pine germination and seedling survival may vary greatly across its geographical range, as it is widely distributed within north latitudes. Mediterranean region is a marginal sector within this species range and its ecological conditions differ greatly from those of the bulk of the area. Mortality rates in Mediterranean environments seem to be related to summer weather (mainly drought) and high livestock stocking rather than to cold, light or flooding. Most available information on scots pine regeneration process comes from north European experiences and is not transferable to Spanish forests, whereas studies on Mediterranean region are much scarcer. The present work aims at broadening Scots pine regeneration knowledge within Mediterranean region by analyzing its establishment process in the “Cabeza de Hierro” forest: a Scots pine uneven-aged forest at blocklevel scale, exemplary managed for multi-services purpose. Germination and surviving processes are to be characterized and quantified as to vegetation cover both in trees, shrubs, grass and litter strata. Soil compaction effects on forest sustainability are also assessed and the efficacy of some site preparation techniques on regeneration success is tested and compared (scarification vs. scalping+mounding). Two sampling networks comprising respectively 198 (SN I) and 24 plots (SN II) of 1.5x1.5m have been established over a wide range of vegetal cover conditions within the forest. Soil preparation techniques have been applied only to some of the sampling points; namely, 1 out of 4 plots have been scarified within Sampling Network I , while 1 out of 2 plots have been object of scalping & mounding within Sampling Network II. After localized sowing prior to growing season, germination and surviving have been periodically sampled for either one (SN II) or three years (SN I). Supplementary measures for micrometeorological variables, stand density, ground vegetal cover and compaction have also been carried out. Results obtained for the studied forest lead to the following insights regarding Scots pine natural regeneration process within this sort of forest and environment: 1) seedling establishment success rate is quite low (0,15% of sowing seeds), due to high mortality during the first summer (>92%), following a prior 17% rate of germination over viable seeds reaching the soil; 2) mortality rate remains high until the third year after emergence and then decreases to the 50% of surviving; 3) although vegetal cover significantly affects both seedling germination and survival, lineal correlation between those two processes is rather low, which may indicate that places fit for emergence are not necessarily suitable for summer surviving; 4) soil scarification improves both germination and survival during the first growing season, but it is associated to higher mortality rates during the next two years; hence it has no significant medium term effect; 5) scalping & mounding treatment is more effective than scarification concerning establishment improving during the first summer; but its effects are only significant under intermediate stand density levels; 6) soil compaction does not restrict either forest productivity or persistence, despite the area’s long history of high livestock stocking rates and mechanized logging.
Resumo:
Los análisis de sensibilidad son una herramienta importante para comprender el funcionamiento de los modelos ecológicos, así como para identificar los parámetros más importantes en su funcionamiento. Además, los análisis de sensibilidad pueden utilizarse para diseñar de forma más efectiva planes de muestreo de campo dirigidos a calibrar los modelos ecológicos. En los estudios de ecosistemas forestales, el análisis cuantitativo de la parte subterránea es mucho más costoso y complicado que el estudio de la parte aérea, en especial el estudio de la dinámica de producción y descomposición de raíces gruesas y finas de los árboles. En este trabajo se muestra un ejemplo de análisis de sensibilidad del modelo forestal FORECAST a parámetros que definen la biomasa, longevidad y concentración de nitrógeno en las raíces de los árboles. El modelo se calibró para simular dos rodales de pino silvestre (Pinus sylvestris) en los Pirineos de Navarra. Los resultados indican que la tasa de renovación de raíces finas es el parámetro más influyente en las estimaciones del modelo de crecimiento de los árboles, seguida de la concentración de N en las mismas, siendo la relación biomasa subterránea/total el parámetro al cual el modelo es menos sensible. Además, el modelo es más sensible a los parámetros que definen el componente subterráneo de la biomasa arbórea cuando simula un sitio de menor capacidad productiva y mayor limitación por nutrientes.
Resumo:
We compared within-population variability and degree of population differentiation for neutral genetic markers (RAPDS) and eight quantitative traits in Central American populations of the endangered tree, Cedrela odorata. Whilst population genetic diversity for neutral markers (Shannon index) and quantitative traits (heritability, coefficient of additive genetic variation) were uncorrelated, both marker types revealed strong differentiation between populations from the Atlantic coast of Costa Rica and the rest of the species' distribution. The degree of interpopulation differentiation was higher for RAPD markers (F-ST 0.67 for the sampled Mesoamerican range) than for quantitative traits (Q(ST) = 0.30). Hence, the divergence in quantitative traits was lower than could have been achieved by genetic drift alone, suggesting that balancing selection for similar phenotypes in different populations of this species. Nevertheless, a comparison of pair-wise estimates of population differentiation in neutral genetic markers and quantitative traits revealed a strong positive correlation (r = 0.66) suggesting that, for C. odorata, neutral marker divergence could be used as a surrogate for adaptive gene divergence for conservation planning. The utility of this finding and suggested further work are discussed.
Resumo:
Stylization is a method of ornamental plant use usually applied in urban open space and garden design based on aesthetic consideration. Stylization can be seen as a nature-imitating ornamental plant application which evokes the scenery rather than an ecological plant application which assists the processes and functions observed in the nature. From a different point of view, stylization of natural or semi-natural habitats can sometimes serve as a method for preserving the physiognomy of the plant associations that may be affected by the climate change of the 21st century. The vulnerability of the Hungarian habitats has thus far been examined by the researchers only from the botanical point of view but not in terms of its landscape design value. In Hungary coniferous forests are edaphic and classified on this basis. The General National Habitat Classification System (Á-NÉR) distinguishes calcareous Scots pine forests and acidofrequent coniferous forests. The latter seems to be highly sensitive to climate change according to ecological models. The physiognomy and species pool of its subtypes are strongly determined by the dominant coniferous species that can be Norway spruce (Picea abies) or Scots pine (Pinus sylvestris). We are going to discuss the methodology of stylization of climate sensitive habitats and briefly refer to acidofrequent coniferous forests as a case study. In the course of stylization those coniferous and deciduous tree species of the studied habitat that are water demanding should be substituted by drought tolerant ones with similar characteristics. A list of the proposed taxa is going to be given.
Resumo:
Drought is a key factor affecting forest ecosystem processes at different spatio-temporal scales. For accurately modeling tree functioning ? and thus for producing reliable simulations of forest dynamics ? the consideration of the variability in the timing and extent of drought effects on tree growth is essential, particularly in strongly seasonal climates such as in the Mediterranean area. Yet, most dynamic vegetation models (DVMs) do not include this intra-annual variability of drought effects on tree growth. We present a novel approach for linking tree-ring data to drought simulations in DVMs. A modified forward model of tree-ring width (VS-Lite) was used to estimate seasonal- and site-specific growth responses to drought of Scots pine (Pinus sylvestris L.), which were subsequently implemented in the DVM ForClim. Ring-width data from sixteen sites along a moisture gradient from Central Spain to the Swiss Alps, including the dry inner Alpine valleys, were used to calibrate the forward ring-width model, and inventory data from managed Scots pine stands were used to evaluate ForClim performance. The modified VS-Lite accurately estimated the year-to-year variability in ring-width indices and produced realistic intra-annual growth responses to soil drought, showing a stronger relationship between growth and drought in spring than in the other seasons and thus capturing the strategy of Scots pine to cope with drought. The ForClim version including seasonal variability in growth responses to drought showed improved predictions of stand basal area and stem number, indicating the need to consider intra-annual differences in climate-growth relationships in DVMs when simulating forest dynamics. Forward modeling of ring-width growth may be a powerful tool to calibrate growth functions in DVMs that aim to simulate forest properties in across multiple environments at large spatial scales.
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In this study, we investigated the relationship between vegetation and modern-pollen rain along the elevational gradient of Mount Paggeo. We apply multivariate data analysis to assess the relationship between vegetation and modern-pollen rain and quantify the representativeness of forest zones. This study represents the first statistical analysis of pollen-vegetation relationship along an elevational gradient in Greece. Hence, this paper improves confidence in interpretation of palynological records from north-eastern Greece and may refine past climate reconstructions for a more accurate comparison of data and modelling. Numerical classification and ordination were performed on pollen data to assess differences among plant communities that beech (Fagus sylvatica) dominates or co-dominates. The results show a strong relationship between altitude, arboreal cover, human impact and variations in pollen and nonpollen palynomorph taxa percentages.
Resumo:
How rainfall infiltration rate and soil hydrological characteristics develop over time under forests of different ages in temperate regions is poorly understood. In this study, infiltration rate and soil hydrological characteristics were investigated under forests of different ages and under grassland. Soil hydraulic characteristics were measured at different scales under a 250-year-old grazed grassland (GL), 6-year-old (6yr) and 48-year-old (48yr) Scots pine (Pinus sylvestris) plantations, remnant 300-year-old individual Scots pine (OT) and a 4000-year-old Caledonian Forest (AF). In situ field-saturated hydraulic conductivity (Kfs) was measured, and visible root:soil area was estimated from soil pits. Macroporosity, pore structure and macropore connectivity were estimated from X-ray tomography of soil cores, and from water-release characteristics. At all scales, the median values for Kfs, root fraction, macroporosity and connectivity values tended to AF>OT>48yr>GL>6yr, indicating that infiltration rates and water storage increased with forest age. The remnant Caledonian Forest had a huge range of Kfs (12 to >4922mmh-1), with maximum Kfs values 7 to 15 times larger than those of 48-year-old Scots pine plantation, suggesting that undisturbed old forests, with high rainfall and minimal evapotranspiration in winter, may act as important areas for water storage and sinks for storm rainfall to infiltrate and transport to deeper soil layers via preferential flow. The importance of the development of soil hydrological characteristics under different aged forests is discussed.
Resumo:
Little is known about historic wood as it ages naturally. Instead, most studies focus on biological decay, as it is often assumed that wood remains otherwise stable with age. This PhD project was organised by Historic Scotland and the University of Glasgow to investigate the natural chemical and physical aging of wood. The natural aging of wood was a concern for Historic Scotland as traditional timber replacement is the standard form of repair used in wooden cultural heritage; replacing rotten timber with new timber of the same species. The project was set up to look at what differences could exist both chemically and physically between old and new wood, which could put unforeseen stress on the joint between them. Through Historic Scotland it was possible to work with genuine historic wood from two species, Oak and Scots pine, both from the 1500’s, rather than relying on artificial aging. Artificial aging of wood is still a debated topic, with consideration given to whether it is truly mimicking the aging process or just damaging the wood cells. The chemical stability of wood was investigated using Fourier-transform infrared (FTIR) microscopy, as well as wet chemistry methods including a test for soluble sugars from the possible breakdown of the wood polymers. The physical properties assessed included using a tensile testing machine to uncover possible differences in mechanical properties. An environmental chamber was used to test the reaction to moisture of wood of different ages, as moisture is the most damaging aspect of the environment to wooden cultural objects. The project uncovered several differences, both physical and chemical, between the modern and historic wood which could affect the success of traditional ‘like for like’ repairs. Both oak and pine lost acetyl groups, over historic time, from their hemicellulose polymers. This chemical reaction releases acetic acid, which had no effect on the historic oak but was associated with reduced stiffness in historic pine, probably due to degradation of the hemicellulose polymers by acid hydrolysis. The stiffness of historic oak and pine was also reduced by decay. Visible pest decay led to loss of wood density but there was evidence that fungal decay, extending beyond what was visible, degraded the S2 layer of the pine cell walls, reducing the stiffness of the wood by depleting the cellulose microfibrils most aligned with the grain. Fungal decay of polysaccharides in pine wood left behind sugars that attracted increased levels of moisture. The degradation of essential polymers in the wood structure due to age had different impacts on the two species of wood, and raised questions concerning both the mechanism of aging of wood and the ways in which traditional repairs are implemented, especially in Scots pine. These repairs need to be done with more care and precision, especially in choosing new timber to match the old. Within this project a quantitative method of measuring the microfibril angle (MFA) of wood using polarised Fourier transform infrared (FTIR) microscopy has been developed, allowing the MFA of both new and historic pine to be measured. This provides some of the information needed for a more specific match when selecting replacement timbers for historic buildings.
Resumo:
The purpose of this work in progress study was to test the concept of recognising plants using images acquired by image sensors in a controlled noise-free environment. The presence of vegetation on railway trackbeds and embankments presents potential problems. Woody plants (e.g. Scots pine, Norway spruce and birch) often establish themselves on railway trackbeds. This may cause problems because legal herbicides are not effective in controlling them; this is particularly the case for conifers. Thus, if maintenance administrators knew the spatial position of plants along the railway system, it may be feasible to mechanically harvest them. Primary data were collected outdoors comprising around 700 leaves and conifer seedlings from 11 species. These were then photographed in a laboratory environment. In order to classify the species in the acquired image set, a machine learning approach known as Bag-of-Features (BoF) was chosen. Irrespective of the chosen type of feature extraction and classifier, the ability to classify a previously unseen plant correctly was greater than 85%. The maintenance planning of vegetation control could be improved if plants were recognised and localised. It may be feasible to mechanically harvest them (in particular, woody plants). In addition, listed endangered species growing on the trackbeds can be avoided. Both cases are likely to reduce the amount of herbicides, which often is in the interest of public opinion. Bearing in mind that natural objects like plants are often more heterogeneous within their own class rather than outside it, the results do indeed present a stable classification performance, which is a sound prerequisite in order to later take the next step to include a natural background. Where relevant, species can also be listed under the Endangered Species Act.
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Growing evidence suggests environmental change to be most severe across the semi-arid subtropics, with past, present and projected drying of the Mediterranean Basin posing a key multidisciplinary challenge. Consideration of a single climatic factor, however, often fails to explain spatiotemporal growth dynamics of drought-prone ecosystems. Here, we present annually resolved and absolutely dated ring width measurements of 871 Scots pines (Pinus sylvestris) from 18 individual plot sites in the Central Spanish Pinar Grande forest reserve. Although comprising tree ages from 6 to 175 years, this network correlates surprisingly well with the inverse May–July diurnal temperature range (r = 0.84; p < 0.00011956–2011). Ring width extremes were triggered by pressure anomalies of the North Atlantic Oscillation, and the long-term growth decline coincided with Iberian-wide drying since the mid-1970s. Climate model simulations not only confirm this negative trend over the last decades but also project drought to continuously increase over the 21st century. Associated ecological effects and socio-economic consequences should be considered to improve adaptation strategies of agricultural and forest management, as well as biodiversity conservation and ecosystem service.
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NADP+-isocitrate dehydrogenase (NADP+-IDH; EC 1.1.1.42) is involved in the supply of 2-oxoglutarate for ammonia assimilation and glutamate synthesis in higher plants through the glutamine synthetase/glutamate synthase (GS/GOGAT) cycle. Only one NADP+-IDH form of cytosolic localization was detected in green cotyledons of pine (Pinus spp.) seedlings. The pine enzyme was purified and exhibited molecular and kinetic properties similar to those described for NADP+-IDH from angiosperm, with a higher catalytic efficiency (105 m−1 s−1) than the deduced efficiencies for GS and GOGAT in higher plants. A polyclonal antiserum was raised against pine NADP+-IDH and used to assess protein expression in the seedlings. Steady-state levels of NADP+-IDH were coordinated with GS during seed germination and were associated with GS/GOGAT enzymes during chloroplast biogenesis, suggesting that NADP+-IDH is involved in the provision of carbon skeletons for the synthesis of nitrogen-containing molecules. However, a noncoordinated pattern of NADP+-IDH and GS/GOGAT was observed in advanced stages of cotyledon development and in the hypocotyl. A detailed analysis in hypocotyl sections revealed that NADP+-IDH abundance was inversely correlated with the presence of GS, GOGAT, and ribulose-1,5-bisphosphate carboxylase/oxygenase but was associated with the differentiation of the organ. These results cannot be explained by the accepted role of the enzyme in nitrogen assimilation and strongly suggest that NADP+-IDH may have other, as-yet-unknown, biological functions.