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     

Variation with Tree Age of Needle Cuticle Topography and Stomatal Structure in Pinus radiate D. Don

Scanning electron microscopy reveals differences in the surfacetopography and stomatal structure of 1-year-old needles of Pinusradiata sampled from trees of different ages. The cuticularridges on young-tree needles show an even pitch, whereas theridges on mature-tree needles appear slightly puckered, withsmall discontinuities. The stomata on mature-tree needles havea smaller pore (10–15 µm) than young-tree needles(15–20 µm). In young trees a fine rodlet, or tubularwax covers the walls of the guard and subsidiary cells. Thestomatal antechamber predominating in mature-tree needles containsan amorphous wax, which frequently closes the pore between theoverarching stomatal lips. The yield of crude wax from chloroformextracts of needles of trees of all ages is approximately 0.2per cent, and there is more of the acidic component in the waxof mature-tree needles. It is suggested that wax occlusionswithin the stomatal antechamber of P. radiata may contributeto mature-tree resistance to the needle pathogen, Dothistromapini Hulbary.     

马尾松毛虫对马尾松和湿地松的选择与适应研究

用马尾松老叶、新叶,湿地松老叶、新叶,当年生马尾松新叶、湿地松新叶和当年生以20％和40％β-蒎烯处理过的马尾松新叶作食料饲养马尾松毛虫。结果表明：马尾松毛虫幼虫在第1代发生期取食马尾松老叶、马尾松新叶、湿地松老叶和湿地松新叶等4种不同质量的食料后,因其食料中10种必需氨基酸含量和挥发性物质β-蒎烯含量的不同,导致幼虫的存活率差异显著。其中,取食马尾松老叶的幼虫成活率最高（44％）,依次为马尾松新叶（32.7％）、湿地松老叶（4％）,而取食湿地松新叶的幼虫全部死亡。通过马尾松针叶增加外援β-蒎烯喂养马尾松毛虫实验,表明增加了外源β-蒎烯的松针饲养的马尾松毛虫幼虫与用同样松针未增加外源β-蒎烯饲养的幼虫相比,死亡率明显提高;松树中β-蒎烯含量的增加与幼虫成活率呈显著负相关。由此,我们阐明了马尾松与湿地松对马尾松毛虫的抗虫机制。     

Growth and Nutrient Retranslocation in Needles of Radiata Pine in Relation to Nitrogen Supply

The effects of N application on tree growth and the retranslocationof N, P, and K from young needles to new growth were examinedin young radiata pine (Pinus radiata D. Don) trees. Nitrogen fertilization increased the number and size of needles,rates of shoot production, stem volume growth and tree biomass.Foliar N and P contents (µg per needle) fluctuated ina cyclic fashion with prominent phases of accumulation, retranslocationand replenishment. The patterns of these fluctuations in controland N-fertilized trees were similar, although the fluxes ofN, P and K in and out of needles were increased by N fertilization.Greater translocation (g per tree) of N and K from needles ofN fertilized trees occurred because fertilization increasedthe needle weight and the proportion of N and K retranslocatedfrom individual needles. Nitrogen fertilization increased theretranslocation of P largely as a result of higher needle mass.Trees supplied with more than adequate amounts of P in the soilretranslocated up to 58 per cent of the initial pool of P fromyoung needles. The periods of high retranslocation coincidedwith periods of high concentrations of soil mineral N and withshoot production. Conversely, the periods of rapid replenishmentof N and P into the needles coincided with the time of slowshoot growth and low concentration of soil mineral N. The growthrate of trees, rather than the availability of nutrients inthe soil was the main factor controlling retranslocation. For radiata pine, retranslocation from needles is not a mechanismspecific for coping with low soil fertility. It seems to bea mechanism which enhances the nutrient supply to apical growingpoints, especially during periods of flushing. Pinus radiata, nitrogen supply, shoot growth, nutrient fluctuations and retranslocation, nutrient use and adaptation     

Within-crown Foliar Plasticity of Western Hemlock, Tsuga heterophylla, in Relation to Stand Age

Anatomy and morphology of needles from six different positionswithin the crowns of western hemlock,Tsuga heterophylla (Raf.)Sarg., were studied to investigate the plastic response of leafstructure to crown position, in relation to stand age. Treeswere sampled from three stands, all of which originated followingcatastrophic fires, representing a chronosequence (15, 55 and145 years old) of stand development. Needles from the highestand outermost crown positions showed morphological and anatomicalresponses normally associated with ‘sun leaves’.As a measure of expressed plasticity, we calculated a relativetrait range (RTR) index to quantify differences in expressedtraits between needles from the upper outer crown (UO needles)and those from the lower inner crown (LI needles). RTR was positivefor most morphological and anatomical traits, indicating largertrait measurements on UO needles relative to LI needles (e.g.UO needles were thicker and had more vascular tissue than LIneedles). However, the degree to which sun and shade leaf traitswere expressed varied between stands. In the youngest stand,UO needles had higher maximal rates of photosynthesis than LIneedles (positive RTR_Ps), but this trend was reversed in theoldest stand (negative RTR_Ps). Mean RTR across all morphologicaland anatomical traits was higher in the 145-year-old stand thanin either the 55-year-old stand (P = 0·03) or the 15-year-oldstand (P < 0·01). While this probably relates mainlyto differences in light conditions between the three stands,it may also indicate a connection between ontogeny and phenotypicplasticity. Copyright 2001 Annals of Botany Company Foliar plasticity, needle anatomy and morphology, ontogeny, photosynthesis, stand development, sun–shade, Tsuga heterophylla(Raf.) Sarg., western hemlock     

The presence of structural surface waxes on coniferousneedles affects the pattern of dry deposition of fine particles

Monodisperse particles (particle diameter 0.5 µm) wereproduced by a particle generator and tagged witha fluorescentdye. The particles were injected into asmall wind tunnel, intowhich single needles or small branches of Picea abies, Pinussylvestris, and Abies alba had been introduced. The needleswere examined after treatment, using fluorescence microscopy,and the spatial patterns of particle deposition determined. The particles deposited preferentially in the stomatal regionsof the needles. In these areas the incidence of micro-roughnessdue to epicuticular waxes is highest, reducing the laminar boundarylayer of the needle. Atmospheric particles of less than 1 /mdiameter are mostly hygroscopic and the potential effect ofsubstantial deposition of these particles to stomatal surroundingsand their influence on plant water relations is discussed. Key words: Conifers, air pollution, stomata, dry deposition, aerosols     

Spatial Distribution of Ethylene Production by Individual Needles Along a Shoot ofPinus sylvestrisL.: Relationship with Peroxidase Activity

Spatial distribution of ethylene production by individual currentyear needles and the relationship with peroxidase activity wasstudied along a shoot of Scots pine (Pinus sylvestrisL.). Muchvariation in ethylene production between and within individualseedlings was found. There was a significant difference in meanintensity of ethylene production between orthostichies withinindividual seedlings. The capacity of individual needles toconvert 1-aminocyclopropane-1-carboxylic acid (ACC) to ethylenedid not correlate with the basal rate of ethylene production.Mean ethylene production intensity by orthostichies correlatedpositively with mean peroxidase activity. A positive correlationbetween the intensity of ethylene production by individual needlesand detachment-induced peroxidase activity was found. The datasuggests that the rate of ethylene production by individualneedles may be directly related to their peroxidase activityand at least in part to their antioxidative capacity.Copyright1998 Annals of Botany Company Pinus sylvestris,pine, ethylene production, spatial distribution, individual needles, peroxidase, seasonal changes.     

Electron Microscopy of the Epidermis and Cuticle of the Needles of Pinus nigra var. maritima in relation to infection by Lophodermella sulcigena

Changes in the ultrastructure of the epidermis and cuticle ofCorsican pine (Pinus nigra var. maritima (Aiton) Melville) inrelation to infection of the needle by Lophodermella sulcigena(Rostr.) v Hohn were recorded throughout a growing season. Epidermalcells when young had thin walls and a thin smooth cuticle: bothstructures thickened with age, and the needle surface becamerough. Densely packed wax tubes developed around the stomatalpore and later became scattered over the needle surface. Themicro-flora present on the needles is briefly noted. Changesin the structure and micro-flora of needles as they aged arethought to contribute to the confinement of infection by L.sulcigena to the needle bases.     

Structure of the Needles in the Early Phases of Development in Pinus ponderosa P. et C. Lawson with Special Reference to Plastids

Before they emerged from the fascicular sheath, the tissuesof young needles of Pinus ponderosa P. et C. Lawson alreadyshowed some characteristics typical of mature needles. The organelles,particularly the plastids, had undergone different development.The plastids in different types of cell varied in their ultrastructureand in association with the endoplasmic reticulum (ER). A sheathof ER was observed around the amoeboid plastids in epidermalcells, epithelial cells of resin ducts and maturing transfusiontracheids whereas there was no ER sheath around the young mesophyllchloroplasts, the fusiform chloroplasts in some transfusiontracheids and the proplastids in xylem and phloem cells. Thecontent of chlorophyll (a+b) was 0·85 g kg^-1 dry matterand chlorophyll a/b ratio was 2·70. The needles may becomephotosynthetically active whilst still within the fascicularsheaths.Copyright 1993, 1999 Academic Press Pinus ponderosa, ponderosa pine, needle structure, needle ontogeny, plastids, endoplasmic reticulum     

The uptake of and structural changes induced by trichloroacetic acid in the needles of Scots pine seedlings

The uptake of trichloroacetic acid (TCA) and structural changesinduced in the needles of Scots pine (Pinus sylvestris L.) seedlingswere studied. Two exposure set-ups, a root route and an atmosphericroute through the surfaces simulating the wet deposition offog, were used. Both set-ups included two dose levels and correspondingcontrol treatments. The temperature and the relative humidityin the climate chambers were adjusted to represent the conditionsof June–July in a subarctic area in central Finland. Theseedlings were exposed three times a week for two months. Theresults showed that the uptake of TCA in needles occurred bothvia roots and via needle surface. However, most of the TCA viathe atmospheric route was absorbed on the surface of the needles.The structural responses in pine needles depended partly onthe treatment method: TCA applied via the atmospheric routedisintegrated the structures of the epicuticular waxes and thatof the stomatal cells, which was not seen in the exposures viaroots. A common feature was the decrease in size of the chloroplastsin concert with the increasing TCA concentrations inside theneedles. Key words: Pinus sylvestris L., climate chamber, effects, microscopic structure, secondary pollutant, TCA, trichloroacetic acid     

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     

Long-distance Transport of Reduced Sulphur in Spruce (Picea abies L.)

Individual needles of 4-year-old spruce trees (Picea abies L.)were exposed to ³⁵S-GSH via the cut surface of the needle tipfor 2–3 h at different times during the growing season.From the previous year's source needle, ³⁵S-GSH was exportedinto the twig and translocated predominantly towards the twigapex. Independent from the growing season, transport towardsthe stem did not exceed 20% of the radioactivity recovered fromthe tissue samples. In July, the new needles appeared to bemetabolic sinks for reduced sulphur as indicated by the higherpercentages of both soluble and insoluble ³⁵S in this twig section,as compared to the previous years' needles. In October, thenew needles were no longer found to be a preferential sink forreduced sulphur. Estimates of the rate of GSH export from thesource needle in the light (2.5–11.3 nmol GSH g     

Dry Weight and N Partitioning in Relation to Substrate N Supply, Internal N Status and Developmental Stage in Jack Pine (Pinus banksianaLamb.) Seedlings: Implications for Modelling

Dry weight and nitrogen (N) partitioning of sand-cultured youngjack pine (Pinus banksianaLamb.) seedlings under controlledenvironments were studied 3, 6, 9, 12 and/or 15 weeks afterthe initiation of six dynamic N supply treatments. The supplyof other nutrients was maintained at optimal levels. Total dryweight varied widely among treatments and whole plant totalN concentration ranged from 10 to 32 mg g^-1d. wt at most samplingintervals. Whole plant N concentration changed, with time, accordingto three distinct patterns: (1) stable; (2) rapidly increasing;or (3) gradually declining. Regardless of N treatment and samplinginterval, whole plant N concentration was linearly and positivelycorrelated with root, needle and stem N concentration. Dry weightand N weight ratios of needles declined, whereas those of rootsincreased linearly with decreasing whole plant N concentration(r²=0.43 to 0.76) regardless of N regime. Dry matter partitioningto stems, however, was better explained by developmental stagethan by whole plant N concentration. With the decline in internalN status, N was increasingly concentrated in roots at the expenseof needles and stems. These results suggest: (1) dry weightand N partioning may be largely a function of the internal Nstatus of plant rather than root and shoot activities; (2) bothshoot and root specific activities may have a close, positiveassociation with whole plant N concentration; (3) N-partitioningmay be an active process itself and may warrant separate considerationfrom dry weight; and (4) developmental stage may be a significantdeterminant of partitioning, particularly to stems.Copyright1998 Annals of Botany Company Developmental stage; dry weight; internal N status; jack pine; modelling; nitrogen; partitioning;Pinus banksiana; substrate N supply.     

Seasonal Changes in Respiration, Photosynthesis, and Translocation of the 14C Labelled Products of Photosynthesis in Young Pinus strobus L. Plants

Three-year-old Pinus strobus plants, grown under conditionsof either high or low light intensities, were brought from thenursery to the laboratory every three to four weeks from themiddle of April 1961 until January 1962. Translocation, measuredas the amount of ¹⁴C recovered from the roots at the end ofseven hours of illumination following exposure of the shootto ¹⁴CO₂, was found to be high in the spring, dropping to negligibleamounts during June and July, increasing again in the autumnand declining after October. Seasonal variation in root respirationwas found to parallel that of translocation. Rates of apparentphotosynthesis were low during the spring, rising to a maximumduring September, and then declining over the winter. The respiration,photosynthesis, and translocation of the low-light grown plantsfollowed a similar pattern to those grown in high-light, exceptthat in general rates were of a lower order. In the high-light grown plants more than 90 per cent of theabsorbed carbon was present in the ethanol-soluble form, ofwhich sugars formed at least 90 per cent. This was even morepronounced in the case of low-light grown plants. The main sugarwas always sucrose. The raffinose content was found to decreaseduring the warmer months. The new needles, during their period of maximum growth, fixedcarbon dioxide photosynthetically at a rate comparable to thatof the old needles. The new stems also possessed a relativelyhigh carbon dioxide fixing ability. Shoot growth, as measured by the increase in length of the newleader stem and new needles, showed the typical patterns forpine species.     

Water relations, xylem embolism and histological features of Pinus thunbergii inoculated with virulent or avirulent pine wood nematode, Bursaphelenchus xylophilus

The development process of pine wilt disease caused by Bursaphelenchusxylophilus (Steiner and Buhrer) Nickle, pine wood nematode,was studied ecophysiologically and histologically in relationto pathogenicity of B. xylophilus. Judging from the predawnxylem pressure potential of needles, the heat pulse velocity,and the soil water potential, the control Pinus thunbergii Parl.,used for the study, was not water-stressed. Virulent B. xylophilusisolate can kill non-waterstressed pines. In virulent B. xylophilusisolateinoculated pines, the predawn xylem pressure potentialof needles abruptly decreased when the colour of 1-year-oldneedles changed to brown and then the water conducting functionof the xylem was lost completely. Avirulent B. xylophilus isolatedoes not affect the needle colour and the xylem pressure potentialof pines. Avirulent B. xylophilus isolate-inoculated pines,however, responded to nematode invasion by decreasing hydraulicconductance of stem and root xylems. In addition, oleoresinexudation slightly decreased. The decreased hydraulic conductanceresults from embolism of tracheids caused by cavitation in thecentral part of the xylem. From histological observation, allof the parenchyma cells in virulent B. xylophilus isolateinoculatedpines died. In contrast, the parenchyma cells, degenerated inavirulent B. xylophilus isolateinoculated pines, were limitedin the embolized region of the xylem. The difference betweenthe response of pine to the virulent B. xylophilus isolate invasionand that to avirulent B. xylophilus isolate invasion indicatesthat nematode-induced death of pine relates to the death ofparenchyma cells, as well as the decrease in xylem hydraulicconductance. Key words: Embolism, hydraulic conductance, parenchyma cells, pathogenicity of pine wood nematode, pine wilt disease.     

Effects of drought on photosynthesis and on the thermotolerance of photosystem II in seedlings of cedar (Cedrus atlantica and C. libani)

The rate of net CO₂ assimilation (A) in cedar needles declinedduring drought progression, while the quantum yield of electrontransport measured under moderate light and high CO₂ (     

Sap Exudation via the Epidermis of Leaves

Reports of stomatal guttation in Gladiolus and of active waterexudation in Vidafaba could not be confirmed by an experimentaland anatomical investigation. Exudation of water from detachedpine needles is considered to be due to processes distinct fromthose causing guttation. Water soaking of leaves, or the flooding of leaf air spaces,and the wettability of cell walls under root pressure as distinctfrom their wettability by water externally applied are discussedin relation to the experimental results.     

Characterisation of Acidic and Basic Apoplastic Peroxidases from Needles of Norway Spruce (Picea abies, L., Karsten) with Respect to Lignifying Substrates

Acidic and basic peroxidases, termed as POD-A and POD-B, wereisolated from the apoplastic space of spruce (Picea abies, L.)needles and purified by acetone precipitation and anion exchangechromatography to apparent homogeneity. The molecular massesof POD-A and POD-B were 39.6 and 29.0 kDa, respectively. ThepH optimum of both isozymes ranged from 4.5 to 6. The apparentK_m values of POD-A and POD-B were 460 and 210 µM for coniferylalcohol. Both isozymes acted also as NADH oxidases with apparentK_m-values of 103 µM (POD-A) and 70 µM (POD-B). NAD⁺but not NADH was found in the apoplastic space of lignifyingneedles. Based on the lignification rate, the contents and kineticproperties of PODs, NADH oxidation by POD is not the major sourceof H₂O₂ required for lignin polymerisation. (Received December 21, 1996; Accepted March 3, 1997)     

Persistent Effects of Short-term, High Exposure to Chlorine Gas on Physiology and Growth of Pinus ponderosa andPseudotsuga menziesii

Following a single acute exposure to chlorine gas, persistenteffects on epicuticular waxes, cuticular transpiration, treegrowth and mortality were studied in foliage of Pinus ponderosaand Pseudotsuga menziesii for three growing seasons. Chlorinegas exposure caused foliar injury to both exposed foliage andfoliage that flushed after exposure (P < 0.05). The tendencyto form films of water rather than droplets was greater in directlyexposed foliage (P < 0.001). Rates of cuticular transpirationwere higher for directly and indirectly exposed foliage of Pinusponderosa up to 1 year after exposure and up to 6 months afterexposure for directly exposed Pseudotsuga menziesii(P < 0.001),after which P. menziesii needles defoliated. Total water content(TWC) and relative water content were significantly correlatedwith foliar injury (P < 0.05). TWC was lower for directlyexposed foliage up to 1 year after exposure (P < 0.001).There was no persistent negative effect on F_v/F_m ratios after1 year. Exposure to chlorine gas did not affect needle lengthor annual shoot increment growth, but exposure was correlatedwith increased bud production. Needle longevity of foliage thatflushed 2 months after exposure was reduced significantly (P< 0.001). Annual stem increment growth for both species decreasedover at least three growing seasons following chlorine gas exposure(P < 0.001), and depended on distance from the spill site.Cone production was lower for exposed Pinus ponderosa treescompared to controls (P < 0.05), and tree mortality was higherwithin approx. 50 m of the release site forPseudotsuga menziesii. Growth responses for both conifers agreed well with predictedpatterns of carbon allocation after defoliation caused by chlorinegas exposure. Copyright 2001 Annals of Botany Company Pinus ponderosa, Pseudotsuga menziesii, conifers, chlorine gas, leaf wettability, cuticular transpiration, water relations, growth, mortality