Root growth potential, seedling morphology and bud dormancy correlate with survival of loblolly pine seedlings planted in December in Alabama

First-year survival of December-planted loblolly pine seedlings sampled from 20 nurseries ranged from 36 to 86%. Various characteristics of the seedlings including root growth potential (RGP), shoot/root ratio, root weight and stem length were correlated with survival. A regression model using the number of new roots >/= 0.5 cm and the shoot/root ratio accounted for 80% of the variation in first-year survival. Alone, the number of new roots >/= 0.5 cm accounted for 76% of the variation in survival. Foliar nutrients were not correlated with survival. Seedlings with a high proportion of quiescent buds at planting had a higher RGP and better survival than seedlings with dormant (endodormant) buds. The data suggest that seedlings with dormant buds should not be planted on sites where rapid new root growth is needed for survival.     

Effects of soil temperature on biomass and carbohydrate allocation in Scots pine (Pinus sylvestris) seedlings at the beginning of the growing season

We studied effects of soil temperature on shoot and root extension growth and biomass and carbohydrate allocation in Scots pine (Pinus sylvestris L.) seedlings at the beginning of the growing season. One-year-old Scots pine seedlings were grown for 9 weeks at soil temperatures of 5, 9, 13 and 17 degrees C and an air temperature of 17 degrees C. Date of bud burst, and the elongation of shoots and roots were monitored. Biomass of current and previous season roots, stem and needles was determined at 3-week intervals. Starch, sucrose, glucose, fructose, sorbitol and inositol concentrations were determined in all plant parts except new roots. The timing of both bud burst and the onset of root elongation were unaffected by soil temperature. At Week 9, height growth was reduced and root extension growth was much less at a soil temperature of 5 degrees C than at higher soil temperatures. Total seedling biomass was lowest in the 5 degrees C soil temperature treatment and highest in the 13 degrees C treatment, but there was no statistically significant difference in total biomass between seedlings grown at 13 and 17 degrees C. In response to increasing soil temperature, below-ground biomass increased markedly, resulting in a slightly higher allocation of biomass to below-ground parts. Among treatments, root length was greatest at a soil temperature of 17 degrees C. The sugar content of old roots was unaffected by soil temperature, but the sugar content of new needles increased with increasing soil temperature. The starch content of all seedling parts was lowest in seedlings grown at 17 degrees C. Otherwise, soil temperature had no effect on seedling starch content.     

Evaluation of three root growth potential techniques with tree seedlings

Three growing systems commonly used to evaluate root growth potential (RGP) are soil, hydroponic, and aeroponic culture. Aeroponic RGP testing is a relatively new technique that has not been adequately compared to conventional methods. This paper reports the results of an experiment designed to compare the amount and variability of root growth of jack pine seedlings in soil, hydroponic, and aeroponic culture. New root length in aeroponic culture was significantly greater at 10, 14, 17, and 21 days and was less variable than that in soil and hydroponic culture. However, the patterns of root growth over time in the three systems are strongly related, so the testing methods provide the same diagnostic information. Root zone temperature differed significantly among locations within each system, but the actual differences were small and did not significantly affect root growth.     

Chlorophyll fluorescence,root growth potential,and stomatal conductance as estimates of field performance potential in conifer seedlings

After cold storage, conifer seedlings in British Columbia are tested for field growth potential before planting. We compared three tests of performance potential using container-grown seedlings of Douglas-fir, interior spruce, lodgepole pine, and western larch (14 seedlots total). On several autumn dates, seedlings were lifted and stored at −2°C. The following spring we tested stored seedlings for root growth potential (RGP), chlorophyll fluorescence (CF), and stomatal conductance (Gs), and then planted seedlings in nursery beds. We assessed survival and shoot dry weight (SDW) after one growing season. Performance test results were significantly correlated with each other (r ≥ 0.47) but showed different relationships with field performance, which varied with lift date. The best performance predictor was the sum of CF and RGP (R ² = 0.79 for 78 seedlot by lift-date combinations), which minimized the risk of planting poor seedlings and not planting good seedlings. A sum of 83 for CF (Fv/Fm %) and RGP (new roots >1 cm) provided a threshold above which survival and growth were good. For evergreen conifers, Gs was a good performance predictor, but required extra time to measure leaf area. We recommend a combination of CF and RGP to assess vigor of shoot and root systems before planting. Wolfgang D. Binder––Scientist Emeritus.     

Shoot and root sensitivity of containerized black spruce,white spruce and jack pine seedlings to late fall freezing

Damage to containerized forest seedlings due to freezing can occur in the fall or early winter in Canadian forest nurseries. The following spring, damage to shoots and impairment of growth is observed. The objectives of this experiment were to measure the impact of late fall low temperatures (0° to --30°C) on whole seedlings of the three most common species used for reforestation in Quebec: black spruce (BS), white spruce (WS) and jack pine (JP). Impacts of freezing temperatures on (i) whole seedling and apical bud mortality, (ii) shoot growth and root mortality, (iii) stem electrical resistance, (iv) shoot and root water relations, (v) concentrations of N, P, K, Ca, Mg, and total sugars in shoots were assessed. JP showed the highest rate of whole seedling mortality while WS showed the highest rate of apical bud mortality. JP was the most severely affected: destruction of the root system at low temperatures as well as a reduction of shoot growth and stem diameter and a decrease (more negative) in shoot and root water potential. WS showed a reduction of shoot growth despite no apparent damage to the root system at low temperatures. BS was not affected by temperatures as low as --30°C. Nutrient and sugar concentrations were not affected by low temperature treatments.     

Photosynthesis and root respiration of Pinus sylvestris and picea abies seedlings after different root freezing and storage temperatures

Photosynthetic performance and root respiration were measured for seedlings of Scots pine and Norway spruce under constant conditions in an open gas exchange system in the laboratory. Measurements were carried out after root exposure to ‐20, ‐5 and 0°C and subsequent longtime storage in darkness at +1 or +4°C. Stomatal conductance in relation to net photosynthetic rates was also investigated after the same treatment of seedlings. Root respiration was low for seedlings whose root system had been exposed to ‐20°C, Scots pine showing lower rates than Norway spruce. This was probably an indication of root damage. At least for one provenance of Scots pine, respiration rates were higher for seedlings stored at +1 than at +4°C. Photosynthetic performance was also lowest for seedlings whose roots had been exposed to +20°C compared to higher temperatures, the difference being more clear‐cut for Norway spruce than for Scots pine. Storage at +1 gave slightly higher photosynthetic rates than at +4°C. There was a close relation between stomatal conductance measured on individual needles and photosynthetic performance measured on the whole seedling.     

Carbon and phosphorus partitioning in Pinus serotina seedlings growing under hypoxic and low-phosphorus conditions

Ten-week-old pond pine (Pinus serotina Michx.) seedlings were grown in solution culture at 5 or 100 microM P and under aerobic or hypoxic solution conditions. After 6 and 10 weeks in the treatments, changes in relative growth rate (RGR), P acquisition and allocation, and carbohydrate partitioning were determined by analyzing tissue for total P, soluble sugars and starch. Six weeks of low-P growth conditions decreased seedling dry weight and the ratio of shoot dry weight to root dry weight (S/R) by 39 and 51%, respectively, in comparison to seedlings from the aerobic, high-P (control) treatment. Mean RGRs of shoots in the low-P treatment were reduced by 33%, whereas root growth was unaffected. After 10 weeks of low-P growth conditions, however, both shoot and root RGRs were significantly reduced, and plants had lower S/R ratios than in any other treatment. Slowed shoot growth was accompanied by starch and nonstructural carbohydrate accumulation in needles, indicating that needle growth was not limited by carbohydrate supply. Six weeks of low-P growth conditions decreased total seedling P by 75%, reflecting a 97% reduction in the net uptake rate (NUR). Shoot NUR as a fraction of seedling NUR was also greatly reduced in the low-P treatment, indicating that low-P growth conditions affected P translocation to the shoot more than P accumulation by roots. In contrast, 6 weeks of hypoxic growth conditions decreased total dry weight of seedlings in the high-P treatment by 41% relative to their aerobic counterparts. Root growth was affected more than shoot growth, however, and S/R ratios increased. After 10 weeks, S/R ratios doubled, primarily because of the reduction in root RGR. Nevertheless, roots of hypoxic seedlings contained a higher percentage of total seedling P than their aerobic counterparts. Net P acquisition per seedling decreased by more than 50% under hypoxic growth conditions, as a result of reductions in both root RGR and seedling NUR. Starch accumulation in shoots of hypoxic seedlings reflected reductions both in root growth and in transport of carbohydrates to nonwoody roots. Carbohydrate availability did not appear to be limiting growth of hypoxic woody roots, which are well-aerated internally, but it may have limited metabolic processes in nonwoody roots of seedlings from the high-P treatment.     

Seedling shoot,needle and bud development in three provenances of Pinus sylvestris and Pinus contorta cultivated in northern Sweden

The patterns of current‐year shoot, needle and terminal bud elongation in seedlings of three Scots pine (Pinus sylvestris L.) and three lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) provenances were compared during the third and fourth growing seasons after planting. Lodgepole pine produced longer shoots and buds than did Scots pine, mainly because lodgepole pine formed more stem units and elongated at a faster rate. Stem unit length and the duration of shoot and bud elongation differed relatively little between species and provenances. Lammas or polycyclic growth occurred in some lodgepole pine provenances, but not in any Scots pine provenance, and was associated with enhanced shoot elongation. Needle elongation commenced earlier, proceeded at a faster rate, and was greater in lodgepole pine than in Scots pine, but ceased about the same time in all species and provenances. The heat sum required to attain 50% of final length was lower for shoots and needles in lodgepole pine than in Scots pine, and for shoots in northern provenances than in southern ones. Mitotic activity in the apical meristem of the terminal bud, which occurred less than one week after the seedlings were free from snow, started and ceased about the same time in each species, but was higher in lodgepole pine than in Scots pine early in the shoot elongation period.     

Genetic and environmental control of seasonal carbohydrate dynamics in trees of diverse Pinus sylvestris populations

We explored environmental and genetic factors affecting seasonal dynamics of starch and soluble nonstructural carbohydrates in needle and twig cohorts and roots of Scots pine (Pinus sylvestris L.) trees of six populations originating between 49 degrees and 60 degrees N, and grown under common garden conditions in western Poland. Trees of each population were sampled once or twice per month over a 3-year period from age 15 to 17 years. Based on similarity in starch concentration patterns in needles, two distinct groups of populations were identified; one comprised northern populations from Sweden and Russia (59-60 degrees N), and another comprised central European populations from Latvia, Poland, Germany and France (49-56 degrees N). Needle starch concentrations of northern populations started to decline in late spring and reached minimum values earlier than those of central populations. For all populations, starch accumulation in spring started when minimum air temperature permanently exceeded 0 degrees C. Starch accumulation peaked before bud break and was highest in 1-year-old needles, averaging 9-13% of dry mass. Soluble carbohydrate concentrations were lowest in spring and summer and highest in autumn and winter. There were no differences among populations in seasonal pattern of soluble carbohydrate concentrations. Averaged across all populations, needle soluble carbohydrate concentrations increased from about 4% of needle dry mass in developing current-year needles, to about 9% in 1- and 2-year-old needles. Root carbohydrate concentration exhibited a bimodal pattern with peaks in spring and autumn. Northern populations had higher concentrations of fine-root starch in spring and autumn than central populations. Late-summer carbohydrate accumulation in roots started only after depletion of starch in needles and woody shoots. We conclude that Scots pine carbohydrate dynamics depend partially on inherited properties that are probably related to phenology of root and shoot growth.     

Early root morphology of jack pine seedlings grown in different types of container

Abstract

This study compared the effects of container type on early root system morphology of jack pine (Pinus banksiana Lamb.) seedlings to determine impacts of container type on root characteristics that may be important for tree stability. Seedlings were grown for one season in Multipots®, Ventblocks®, Copperblocks®, Starpots® and Jiffy® pellets, and for a second season in sand culture in pots. After the first growing season, roots that had grown between Jiffy plugs were cut using a knife, either “early”, in September, or “late”, in November. Seedlings produced in non-pruning containers, Multipots and Ventblocks, had more vertical structural roots, less even root distribution and high shoot:root mass after a second season. Seedlings grown in pruning containers, Copperblocks, Starpots and Jiffy pellets, had more horizontal structural roots and more evenly distributed roots. Jiffy seedlings had high shoot:root mass after pruning, but ratios decreased to low levels similar to Starpots and Copperblocks after a second season in pots. Early pruning of Jiffy seedlings removed less root mass than late pruning, but total live root mass of early and late pruned seedlings was similar after the potting trial. Pruning containers thus produced structural root forms with more desirable characteristics for tree stability. These characteristics were evident after the first season and persisted after the second season in the potting trial.     

Morphological characteristics associated with tolerance to competition from herbaceous vegetation for seedlings of jack pine,black spruce,and white pine

Tolerance of bareroot and container-grown seedlings of black spruce (Picea mariana (Mill.) B.S.P.), jack pine (Pinus banksiana Lamb.), and eastern white pine (Pinus strobus L.) to competition from herbaceous vegetation was examined in the first five years after planting on a site in the Great Lakes/St. Lawrence forest of Ontario, Canada. Shoot and root morphological characteristics of various stocktypes were measured before planting and correlated with 5-year survival and growth following control and no control of herbaceous vegetation. For black spruce and jack pine, medium-sized bareroot stocktypes had greater relative 5-year stem volume growth in the presence of herbaceous vegetation than did container stock of either species or large bareroot stock of spruce. Relative volume growth was measured as the ratio of the cumulative stem volume increment in the presence of vegetation (Veg) to that in the absence of vegetation (NoVeg), i.e., the Veg:NoVeg ratio. In white pine, the Veg:NoVeg ratio of volume increment of medium container and large bareroot stocktypes exceeded that of small container and medium bareroot stocktypes. In jack pine, root collar diameter at planting and number of first-order lateral roots were positively correlated with 5-year Veg:NoVeg ratio of volume increment. In white pine, the Veg:NoVeg ratio was also positively correlated with root collar diameter at planting and with root volume. In black spruce, the ratio was not related to pre-plant morphology. Thus, for white pine and jack pine, certain pre-plant morphological features may be useful in forecasting the relative ability of different stocktypes to grow under herbaceous competition conditions in the field.     

Relationships among cold hardiness, root growth potential and bud dormancy in three conifers

Greenhouse-cultured, container-grown ponderosa pine (Pinus ponderosa var. scopulorum Engelm.), interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) and Engelmann spruce (Picea engelmannii (Parry) Engelm.) were cold acclimated and deacclimated in growth chambers over 19 weeks. Stem cold hardiness, total new root length at 14 days and days to bud break were measured weekly. Relationships among cold hardiness, root growth potential (RGP) and bud dormancy suggest that cold hardiness, which can be measured quickly, could provide a useful basis for estimating the two other parameters. During cold acclimation, there was a lag period in which stem cold hardiness remained at -15 degrees C and RGP was at a minimum, in all three species. Douglas-fir and Engelmann spruce buds remained fully dormant during this lag period. Ponderosa pine buds had no chilling requirement for the loss of dormancy, and reached quiescence during the lag period. Immediately following the lag period, as stem cold hardiness progressed to -22 degrees C, RGP increased to a high plateau in all three species, and Douglas-fir and Engelmann spruce buds approached quiescence. Cold deacclimation and bud development began immediately on exposure to warm, long days, but RGP remained high until stem cold hardiness returned to approximately -15 degrees C. At bud break, cold hardiness and RGP were at the minimum.     

The Influence of Desiccation, Rough Handling and Cold Storage on the Quality and Establishment of Sitka Spruce Planting Stock

In January 3-year-old Sitka spruce were lifted and treated eithercarefully or roughly. Plants in each treatment were either transferredto the laboratory for testing, or transferred to cold storageat –1°C. In the laboratory, half of the plants ineach treatment were desiccated to shoot water potentials ofless than –2.0 MPa. Plants were then tested for root growthpotential (RGP), bud dormancy, frost hardiness and carbohydratecontent before transplanting at a field site, Additional plantswere removed from the nursery and cold store at approximatelymonthly intervals until late April, and then treated as above.In April, plants in each treatment were transferred to on-sitefield storage (sheughs), for 2 or 4 weeks prior to testing asabove. Carefully handled plants maintained large RGP which was positivelycorrelated with plant water potential, root water content, shootrelative growth rate and field survival. Overall RGP was reduced: 59 per cent by desiccation; 85 percent by rough handling; and 98 per cent by desiccation and roughhandling in combination. On site storage for 4 weeks in April/May reduced RGP. Whereplants produced more than 30 new roots > 1 cm long in RGPtests, field survival exceeded 90 per cent. Cold storage at–1°C maintained RGP, (of carefully handled plants),bud dormancy and shoot frost hardiness at approximately pre-storagevalues. In contrast, total nonstructural carbohydrate contentdeclined in cold store but increased in March/April for plantsin the nursery.     

Relationship between carbohydrate concentration and root growth potential in coniferous seedlings from three climates during cold hardening and dehardening

Greenhouse-cultured, container-grown seedlings of Aleppo pine (Pinus halepensis Mill.), radiata pine (Pinus radiata D. Don), and interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) were cold acclimated and deacclimated in growth chambers over 24 weeks. Needle and root cold hardiness and root growth potential (RGP) were measured weekly. Root, needle and stem analyses for soluble sugars and starch were performed biweekly. In all tissues, there was a close correspondence between cold hardiness and the absolute concentration of soluble sugars, as well as between the increase and decrease in concentration of soluble sugars during cold hardening and dehardening, respectively, supporting the theory that soluble sugars function as cryoprotectants in plant tissues. The magnitude of starch concentration did not parallel the magnitude of the cold hardiness attained, and changes in starch concentration were related to production and consumption factors, rather than timing of changes in cold hardiness. The rise and fall of RGP paralleled the rise and fall of total carbohydrate concentration in roots. The behavior of the three species was surprisingly similar, considering the different climates to which they are adapted.     

First-year growth response of cold-stored,nursery-grown aspen planting stock

This research examined the first year growth characteristics of cold stored and transplanted nursery-produced aspen (Populus tremuloides) seedlings (container and bareroot (BR)) and compared it to the growth of seedlings that had not been transplanted (established from germinants in the field) and therefore had an unrestricted root system (UR). Prior to planting, nursery-produced seedlings were placed in cold storage (−3°C) and root growth potential (RGP) and total non-structural carbohydrate (TNC) root reserves were tested at 0, 10, 75 and after 150 (container) and 190 days (BR) of storage. Both container and BR stock had much lower root to shoot ratios (RSRs) and root carbohydrate reserves compared to UR seedlings after 170 days. During storage, root reserves in container stock declined faster than in the BR and UR seedlings. RGP in all nursery stock was the highest after 75 days of storage, while longer storage resulted in shoot dieback and reduced root growth. After the first growing season, UR seedlings were one tenth the size of the nursery stock; however, in the second growing season they had no stem dieback and grew twice the height and stem diameter. The higher RSRs and root reserves in the UR seedlings was likely caused by early bud set in its first year of growth. This suggests that inducing bud set earlier in the growing regime might allow seedlings to increase root mass and carbohydrate reserves.     

Root growth and water use efficiency of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and lodgepole pine (Pinus contorta Dougl.) seedlings

One-year-old Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and lodgepole pine (Pinus contorta Dougl.) seedlings were grown for 17 weeks in 100-cm deep, 7.8-liter containers. Two Douglas-fir provenances, one from a wet and one from a dry site in coastal British Columbia, and two lodgepole pine provenances, one from a wet and one from a dry site in interior British Columbia, were grown in wet (522% water content) or dry (318% water content) peat/vermiculite soil in a factorial design. Each container was sealed so that water loss occurred only through the seedling. Five harvests were made at three to five week intervals and water use, dry matter increment, root length and root weight were determined at each harvest. Stomatal conductance and shoot water potentials were measured during the last 12 weeks of the experiment. Lodgepole pine seedlings had greater dry matter production, water use, stomatal conductance and new root length than Douglas-fir seedlings. New root weight of lodgepole pine seedlings exceeded that of Douglas-fir seedlings during the last five weeks of the experiment, and specific root length (root length per unit root weight) of new roots was higher for lodgepole pine seedlings throughout the experiment. Douglas-fir seedlings showed higher water use efficiency (WUE) than lodgepole pine seedlings, and both species showed higher WUE in the dry soil treatment. Douglas-fir seedlings had lower water potentials and higher water uptake rates per unit of new root length than lodgepole pine seedlings, although water uptake rates per unit of root dry weight showed little difference between species. Soil water treatment influenced specific root length of new roots, water uptake per unit of new root length, and WUE in Douglas-fir seedlings more than in lodgepole pine seedlings.     

Spatial distribution,architecture, and development of the root system of Pinus banksiana Lamb. in natural and planted stands

In the province of Québec, Canada, the majority of planted jack pine (Pinus banksiana Lamb.) seedlings are produced in rigid wall containers. More than 95% of them exhibit deformations of the root system which may induce stem instability. Studies of the root architecture of planted jack pine have been limited to a 30 cm radius from the stem, as barely any studies have been devoted to naturally regenerated stands. Moreover, only a few researches have focused on temporal evolution of root systems. The aim of the present study was to characterize the architectural, spatial, and temporal development of jack pine roots in natural and planted stands. Study sites were located in the continuous boreal forest of Quebec. The plantation was done in 1987, so that the trees were 15 years old at the time of sampling. Trees from natural stand had regenerated after a fire in 1983 and were 13–16 years old. The root systems of 14 jack pine trees per site were manually excavated up to a <5 mm diameter, without regard to their distance from the stem. The number, length, diameter, and the spatial and temporal development of roots were analyzed according to three scales of root architecture: the root system, axes, and segments. Overall, the numbers and lengths of roots were higher with planted pines. However, naturally regenerated trees displayed a better distribution of their roots around the stem and at depth, combined with more rapid length growth during the first years. In natural stands, all the trees had a taproot and 30% of the main roots originated at a depth of more than 20 cm, and they are regularly distributed around the stems. Planted trees did not present a taproot and 97% of the main roots originated in the first 20 cm beneath the soil surface. Moreover, 50% of root length was located in one-third of the area surrounding the stems, an area that corresponded to the furrow. Finally, the annual development of lateral roots in planted stand displayed a 5-year delay when compared with natural stand, which also affected maximum growth length and development of the branching pattern. Root distribution and temporal development are known to play a major role in the stability of aerial parts. Seedling production methods, container type, site preparation and planting techniques need to be examined in greater detail in order to assess their effect throughout the development of the root system. It is necessary to compare different sylvicultural practices and with natural/planted stands to gain a clearer understanding of this problem.     

Carry-over effects of ozone on root growth and carbohydrate concentrations of ponderosa pine seedlings

Ozone exposure decreases belowground carbon allocation and root growth of plants; however, the extent to which these effects persist and the cumulative impact of ozone stress on plant growth are poorly understood. To evaluate the potential for plant compensation, we followed the progression of ozone effects, with particular emphasis on the development of new roots. Ponderosa pine (Pinus ponderosa Dougl. ex Laws.) seedlings were exposed to ozone for 2 years. Following removal of the seedlings from ozone, root growth was assessed to characterize the carry-over effects on new root production, and carbohydrate concentrations were measured to determine if allocation strategies differed among ozone treatments. Four months after removal from ozone, dormant seedlings had significantly lower starch concentrations in stems, coarse roots and fine roots than control seedlings. Following root flushing, starch concentrations in all seedlings decreased, with ozone-treated seedlings containing significantly less starch, sucrose, fructose, glucose and total monosaccharides than control seedlings. There was some evidence that stem starch was mobilized to compensate partially for the lower concentrations of root starch in ozone-treated seedlings; however, there was significantly less new root production in seedlings previously exposed to ozone for 2 years than in control seedlings. Early senescence of older needle age classes, perhaps resulting in inadequate available photosynthate, may be responsible for the reduction in new root production during the year following exposure to ozone. Stored carbohydrate reserves, which were depleted in seedlings previously exposed to ozone, were insufficient to compensate for the ozone-induced reduction in canopy photosynthate. We conclude that there are carry-over effects of ozone exposure on ponderosa pine seedlings, including an enhanced potential for seedling susceptibility to other stresses even in respite years when ozone concentrations are low.     

Effects of exogenous gibberellin and auxin on shoot elongation and vegetative bud development in seedlings of Pinus sylvestris and Picea glauca

The hormonal control of stem unit (foliar appendage and axillary structure, if present, plus subtending internode) number and length was investigated in shoots of Scots pine (Pinus sylvestris L.) and white spruce (Picea glauca (Moench) Voss). Seedlings were treated with six gibberellins (GA1, GA3, GA4, GA5, GA9 and GA20) and two auxins (indole-3-acetic acid (IAA) and naphthaleneacetic acid (NAA)) when either neoformed growth was occurring or the terminal vegetative bud was developing. Hormones were applied by drenching the shoot tip, injecting the stem or spraying the foliage. Combined results for all three application methods indicated that shoot elongation in first-year seedlings (i.e., neoformed growth) was promoted in both species by GA1, GA3, GA4 and, less obviously, by GA9. This promotion was attributable to an increase in length, rather than number, of stem units. However, the number of stem units formed during terminal bud development, as reflected in the number of needles (white spruce) or cataphylls (Scots pine) present on the shoot resulting from the terminal bud, was stimulated by GA1, GA3 and GA4 in both species and by GA9 in Scots pine. The GA-induced increase in the number of preformed stem units was associated with increased bud width in white spruce and increased bud length and resulting shoot length in Scots pine. In contrast, application of IAA or NAA either did not affect or inhibited both neoformed growth and terminal bud stem unit number, depending on the application method and concentration. We conclude that, in the Pinaceae, (1) GA stimulates the activity of both the subapical meristem during neoformed growth and the apical meristem during vegetative bud development, and (2) the early non-hydroxylation pathway, via GA9, is the major route of GA biosynthesis. The role of auxin in the control of stem unit number and length remains to be resolved.     

Effects of root temperature on growth and photosynthesis in conifer seedlings during shoot elongation

Growth and gas exchange characteristics were studied in pine (Pinus sylvestris L.) and spruce (Picea abies Karst.) seedlings grown in hydroponic culture in the presence of N (50 mg l(-1)) and transferred at the start of their second growing season to tap water at 5, 8, 12, 16 or 20 degrees C (air temperature between 18-20 degrees C) for 3 weeks (pine) or 5 weeks (spruce). Root growth of both species was completely inhibited at root temperatures of 5 and 8 degrees C, but increased almost exponentially as root temperature increased. Shoot growth was maximal at 12 degrees C in both pine and spruce and decreased at low root temperatures. In both species, CO(2) uptake was decreased at low root temperatures and appeared to be influenced by the pattern of nitrogen retranslocation. In pine seedlings, as root temperature increased, an increasing proportion of the total nitrogen pool was retranslocated to the new shoot, whereas in spruce seedlings nitrogen was retranslocated to the roots. Differences in the retranslocation of nitrogen in the two species were reflected in the amount of soluble protein in needles, which at the end of the experiment increased with increasing root temperature in pine, but decreased in spruce. Our data suggest that in spruce, but not pine, CO(2) uptake was limited by the amount of Rubisco.