Physiological and morphological responses to nitrogen limitation in jack pine seedlings: potential implications for drought tolerance

The morphological and physiological responses to nitrogen (N) limitation in jack pine (Pinus banksiana Lamb.) seedlings were studied following the initiation of four different dynamic N treatments for six and 15 weeks. The N treatments produced needle N concentrations from 11 to 31 mg g^-1dry weight, and seven-fold difference in dry weight at 15 weeks. Low-N jack pine seedlings: 1) had an higher root/shoot ratio; 2) extended their tap root more rapidly; 3) were better able to maintain turgor when shoot water potential declined; and 4) had a larger dry weight fraction and apoplasmic fraction than seedlings with higher foliar N concentrations. These responses may contribute collectively to enhance drought tolerance in N-limited plants, thereby affecting seedling quality. Modifying nursery fertilization regimes, other than optimal as usually applied, may thus be needed to produce stock for use on particularly droughty sites. Knowledge of the nature of drought at a particular site could be an important consideration when making decisions related to fertilization.     

国外容器苗质量调控技术研究进展

近年来,由于造林重点由宜林地向困难立地转移,对苗木质量的要求也随之增高.文章系统总结近年来国外容器苗的质量调控技术及其对于我国困难立地植被恢复的重要意义,从指数施肥、秋季施肥、底部灌溉、短日照处理和夏季造林、修根技术、容器规格、冻藏和解冻等研究进展进行了论述总结.指数施肥的每次施肥量与植物生长节律和养分需求同步,施肥效果较优.秋季施肥能避免苗木的养分稀释效应,已被应用于很多树种.实施底部灌溉的植物生长不亚于上部灌溉的植物,而底部灌溉用水量少,氮肥淋溶小.夏季造林时,将正处于速生期的苗木进行短日照处理,诱导顶芽形成,促进根系发育,能提高苗木抗性和造林效果.苗木冻藏可使空气中自由态的水凝固成结晶态的冰,苗木感染病菌几率降低,因此冻藏成为针叶树种贮藏的标准方法.在当前研究热点的基础上,论文就未来研究方向进行展望.     

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.     

Soil water content and water relations in planted and naturally regenerated Pinus halepensis Mill. seedlings during the first year in semiarid conditions

The capacity of Aleppo pine (Pinus halpensis Mill.) seedlings to overcome the planting shock in dry conditions was assessed by firstly studying the survival and water status during the first year after planting in relation to soil water content. In spite of receiving only 177 mm rainfall during the year studied, survival of planted Aleppo pine was very high (88.5%). Soil water during summer months (after receiving 67 mm rainfall in winter and spring) was only available at 30–60 cm depth, with tension values of –1.1 and –1.3 MPa in July and August respectively; in these conditions, July predawn xylem water potential measured was –2.5 MPa, and midday potential was –3.6 MPa. According to different authors, these values don't jeopardize the survival of Aleppo pine. In addition, acclimation of outplanted seedlings to environmental conditions was followed by comparing their water relations with those of naturally regenerated seedlings on the site. Predawn and midday xylem water potential showed differences in favour of outplanted seedlings since June, indicating an adjustment to this dry site. Compared to naturally regenerated trees, nursery grown stock of the same age before field planting had much more biomass and higher N and P concentrations and contents; although shoot:root balance and Dickson quality index were not significantly different. Finally, planted seedlings acclimation level during first year was also evaluated by Transplant Stress Index, which value (–0.1278) indicated a slight planting impact.     

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.     

Survival and growth of drought hardened <Emphasis Type="Italic">Eucalyptus pilularis</Emphasis> Sm. seedlings and vegetative cuttings

Forestry requires low mortality of transplanted seedlings. Mortality shortly after planting is often associated with inadequate hydration of transplants. Seedlings can be hardened to the drought conditions they may experience after transplanting by exposing them to controlled drought conditions in the nursery. Eucalyptus pilularis Sm. seedlings were drought hardened by providing nil (severe treatment) or half (mild treatment) the daily irrigation routinely received (control treatment) for up to two non-consecutive days per week during the last 4 weeks of growth in the nursery. Drought hardening reduced stem diameter, seedling leaf area, leaf area per root biomass and seedling quality measured by the Dickson quality index, but increased root:shoot ratio. Hardened seedlings had lower stomatal conductance and leaf water potential on the days they received less irrigation that the control treatment. Hardened seedlings had greater stomatal conductance and were less water stressed than seedlings experiencing drought for the first time indicating hardened seedlings had adjusted physiologically to drought. Survival after transplanting in the controlled drought environment in a glasshouse was enhanced by the hardening treatments. Non hardened seedlings that had had their upper leaves manually removed immediately prior to transplanting to reduce leaf area (top-clipped) had similar survival to hardened seedlings. Stomatal conductance and leaf water potential after transplanting were higher in hardened and top-clipped seedlings than unhardened control seedlings or vegetative cuttings. Survival in the field trial was over 95% for all treatments, possibly as rain fell within 4 days of planting and follow-up rain occurred in the subsequent weeks. Neither the hardened or top-clipped seedlings planted in the field trial had reduced growth, increased propensity to form double leaders or worse stem form than control seedlings when measured at age 3 years.     

Root and shoot allometry of bareroot and container Douglas-fir seedlings

Detailed root and shoot development of bareroot and container Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedling stocktypes were compared during the first growing season after outplanting. The study was installed in raised beds with ideal environmental conditions and at a field reforestation site. Survival at both sites was 98% and did not differ between stocktypes. Seedlings were excavated in spring (5, 8, 12, and 16 weeks after planting) and in fall (35, 40, and 45 weeks after planting). In spring, container seedlings had more numbers of new roots and greater new root and shoot biomass than bareroot seedlings at both sites. In fall, bareroot seedlings consistently averaged more new root growth (though nonsignificant) than container seedlings suggesting that stocktype differences may not continue long-term. Container seedlings had significantly greater water percent than bareroot seedlings at the field site (all sample dates) and the raised bed site (weeks 5, 8, and 40 only). Regardless of environmental conditions or season, seedlings at both sites maintained water percent between 60 and 70% of fresh weight. Seedlings grown in the raised beds had much greater growth than those grown in the field. However, relative growth patterns for the two stocktypes were very similar on each site. The data generated establish baseline differences between stocktypes for root initiation, growth, and allometry during the first year after planting. Challenges associated with root development research are discussed.     

Short-day treatment enhances root egress of summer-planted Picea abies seedlings under dry conditions

Abstract

Second year Norway spruce [Picea abies (L.) Karst.] container seedlings, short-day (SD) treated for 3 weeks in July, were exposed together with untreated control seedlings (Co) to three different drought treatments for 5 weeks after planting in early August. The treatments were: (1) regular watering (0 week drought); (2) 2 weeks of drought and 3 weeks of watering; and (3) no watering (5 week drought). No difference was found in the vigour and shoot xylem water potential between the SD-treated and the Co seedlings after the drought treatments. The root growth decreased less for the SD seedlings than for the Co seedlings along with the increase in the length of the drought period.     

Physiology and morphology of Douglas-fir rooted cuttings compared to seedlings and transplants

Cuttings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) from three open-pollinated families were rooted in two types of tray, and then grown for 1.5 years in a bareroot nursery. During their second winter they were sampled periodically and tested for cold hardiness, dormancy status, root growth potential and various morphological characteristics. Two-year-old seedlings and transplants were tested concurrently for comparison. Rooted cuttings, seedlings and transplants cold hardened at similar rates during early winter, achieving the same level of midwinter hardiness (LT(50) = -18 degrees C) in early January. However, rooted cuttings remained hardier later into the spring than did seedlings or transplants. Rooted cuttings exhibited deeper dormancy in early winter than seedlings or transplants but these differences disappeared after January. Root growth potentials of all three stock types remained above threshold values established for transplants throughout winter. Rooted cuttings had greater stem diameter, higher stem diameter to height ratio, and greater root weight than either seedlings or transplants. This may reflect lower growing densities for the rooted cuttings. Root/shoot ratios of rooted cuttings were greater than for seedlings and similar to those of transplants. Rooted cuttings also had deeper and coarser root systems, which probably reflects lack of wrenching at the nursery.     

Fall fertilization of Pinus resinosa seedlings: nutrient uptake, cold hardiness, and morphological development

? Fall fertilization may increase plant nutrient reserves, yet associated impacts on seedling cold hardiness are relatively unexplored.

? Bareroot red pine (Pinus resinosa Ait.) seedlings in north-central Minnesota, USA were fall fertilized at the end of the first growing season with ammonium nitrate (NH₄NO₃) at 0, 11, 22, 44, or 89 kg N ha^?1. Seedling morphology and cold hardiness [assessed by freeze induced electrolyte leakage (FIEL)] were evaluated six weeks after fertilization and following the second growing season.

? Seedling height and number of needle primordia increased with fertilizer rate for both sampling years. Seedlings fertilized with 44 and 89 kg N ha^?1 attained target height (15 cm) after the second growing season. Shoot and root N concentration increased after the first growing season in fall fertilized seedlings compared to controls. Fall fertilized seedlings had lower FIEL (i.e., increased cold hardiness) compared to controls when tested at ?40 °C after the first growing season, but no significant differences in FIEL of control and fertilized seedlings were observed after the second growing season.

? Results suggest that fall fertilization of red pine seedlings can help render desired target height in the nursery, while maintaining or increasing cold hardiness levels.

     

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.     

Performance and nutrient dynamics of holm oak (<Emphasis Type="Italic">Quercus ilex</Emphasis> L.) seedlings in relation to nursery nutrient loading and post-transplant fertility

Holm oak (Quercus ilex L.) seedlings were exponentially (E) nutrient loaded using incremental increases in fertilizer addition or conventionally (C) fertilized using a constant fertilizer rate during nursery culture. The fertility treatments (mg N plant⁻¹) were control (0), 25E, 100E, and 100C. Subsequently, 1-year-old plants were transplanted under simulated soil fertility gradients in a greenhouse to evaluate effects of nutrient loading and post-transplant fertility on seedling performance. Post-transplant fertility consisted of fertilizing plants at two rates (0 vs. 200 mg N plant⁻¹). A water-soluble fertilizer 20-20-20 was supplied in both nursery and post-transplant experiments. Nutrient loading increased plant N content by 73% in 100E and by 75% in 100C relative to controls, although no significant differences were detected between constant and exponential fertilization regimes at the 100 mg N plant⁻¹ rate. When transplanted, nutrient loading promoted post-transplant root growth relative to shoot, implicating potential to confer competitive advantage to loaded holm oak seedlings after trans-planting. In contrast, post-transplant fertility increased new shoot dry mass by 140% as well as N, P and K content relative to controls. Results suggest that holm oak seedlings can be successfully nutrient loaded in the nursery at higher fertility rates, improving its potential to extend new roots, but alternative fertilization regimes and schedules that better fit nutrient availability to the growth rhythm and conservative strategy of this species must be tested.     

Fall fertilization enhanced nitrogen storage and translocation in Larix olgensis seedlings

Fall nutrient loading of deciduous forest nursery seedlings is of special interest because of foliage abscission and varied translocation patterns. For non-deciduous seedlings in the nursery, fall fertilization typically can reverse nutrient dilution and possibly increase nutrient reserves; however, this technique has received little attention with deciduous conifer trees that translocate nutrients before abscising foliage. We evaluated how fall nitrogen (N) fertilization affected N storage and translocation in the deciduous conifer Olga Bay larch (Larix olgensis Henry) seedlings during the hardening period. Seedlings were supplied with 25 mg N seedling^?1 for 15 weeks before hardening and fall fertilization treatments began with a three week application period of K¹⁵NO₃ at 0, 5, 10 and 15 mg N seedling^?1. During the hardening period, fall N fertilization had little effect on seedling morphology. The N concentration and content of needles decreased dramatically as needles abscised, while that of stems and roots increased. Six weeks after fall N fertilization ceased, all seedlings translocated similar net N from their needles. For the control seedlings, this accounted for 84 % of the N stored in stems and roots. For fall fertilized seedlings, however, the proportion of N stored in stems and roots translocated from needles accounted for only 41–61 % of the total because of absorption of fall fertilizer that was translocated directly to stems and roots. Six weeks after fall fertilization, the distribution pattern of N concentration and content in seedlings was found in this order: stems > fine roots > coarse roots > needles. Our results suggest that providing deciduous conifer seedlings N during hardening, in this case Olga Bay larch, is a way to promote nutrient loading during nursery production.     

Effects of Seedbed Density on One-Year-Old <Emphasis Type="Italic">Fraxinus angustifolia</Emphasis> Seedling Characteristics and Outplanting Performance

To assess the effects of seedling spacing on one-year-old seedling morphology in the nursery, seeds of three provenances of Fraxinus angustifolia were sown at five different seedling spacings within rows of two different spacings in the seedbed. Subsequent growth performance of one-year-old seedlings was assessed by planting in the forest. Within row spacings were: 4.3, 5.0, 6.3, 8.3, and 12.5 cm, and there were five rows at 20 cm apart, or three rows at 33 cm apart across the 1.2 m wide seedbeds. Both spacings within and between rows significantly affected shoot height, root collar diameter, root dry weight and shoot dry weight, but not root/shoot ratio. Wider spacings produced larger seedlings, but only the wider spacing within rows significantly increased fine and coarse root mass. Provenances showed significant differences in diameter, root/shoot ratio, and fine and coarse roots, and they also showed interactions with row spacings in height and diameter measurements. One year after outplanting, diameter growth was significantly related to provenance, and diameter growth was 88% greater for trees from 33 cm nursery row spacing than those from 20 cm nursery row spacing.     

不同水肥处理对海岸松苗木生长的影响

以海岸松1年生苗为材料,研究了不同水分、氮肥供应水平对苗高、地径、茎干重和根干重的影响。结果表明,不同氮肥供应水平对4个指标的影响均达到显著性差异水平,不同水分处理对苗高、地径和茎干重的影响达到显著性差异水平,水肥互作对苗高、茎干重的影响达到显著性差异水平。     

Drought tolerance and transplanting performance of holm oak (Quercus ilex) seedlings after drought hardening in the nursery

Drought stress is the main cause of mortality of holm oak (Quercus ilex L.) seedlings in forest plantations. We therefore assessed if drought hardening, applied in the nursery at the end of the growing season, enhanced the drought tolerance and transplanting performance of holm oak seedlings. Seedlings were subjected to three drought hardening intensities (low, moderate and severe) for 2.5 and 3.5 months, and compared with control seedlings. At the end of the hardening period, water relations, gas exchange and morphological attributes were determined, and survival and growth under mesic and xeric transplanting conditions were assessed. Drought hardening increased drought tolerance primarily by affecting physiological traits, with no effect on shoot/root ratio or specific leaf mass. Drought hardening reduced osmotic potential at saturation and at the turgor loss point, stomatal conductance, residual transpiration (RT) and new root growth capacity (RGC), but enhanced cell membrane stability. Among treated seedlings, the largest response occurred in seedlings subjected to moderate hardening. Severe hardening reduced shoot soluble sugar concentration and increased shoot starch concentration. Increasing the duration of hardening had no effect on water relations but reduced shoot mineral and starch concentrations. Variation in cell membrane stability, RT and RGC were negatively related to osmotic adjustment. Despite differences in drought tolerance, no differences in mortality and relative growth rate were observed between hardening treatments when the seedlings were transplanted under either mesic or xeric conditions.     

Effect of some nursery practices on incidence and severity of diseases,and growth of Eucalyptus grandis seedlings*

Nursery practices influenced incidence and severity of diseases of Eucalyptus grandis viz. damping-off caused by Pythium sp., Rhizoctonia solani and Cylindrocladium quinqueseptatum, web might by R. solani, seedling blight by C. quinqueseptatum and shoot wilt by Sclerotium rolfsii. Growth of seedlings of E. grandis and microclimatic conditions were also affected by nursery practices. Shading with coconut leaf thatch led to low light intensity (av. 1463 lux) with high soil water potential, low soil and ambient temperatures, and high severity of damping-off and web blight diseases and poor shoot:root ratio of seedlings. Seed beds under coirmat had dispersed light (av. 22 299 lux), apparently high severity of seedling blight and shoot wilt, and good growth (snoot:root ratio) of seedlings. In both the types of shading high soil moisture regime and high seed rate contributed to high disease severity as well as low shoot:root ratio of seedlings.     

Effects of two Glomus species on the growth and physiological performance of Sophora davidii seedlings under water stress

Sophora davidii is an important leguminous scrub that is widely used for revegetation in the semiarid Loess Plateau and other arid valley areas of China, where it usually suffers drought stress. This study investigated the effects of arbuscular mycorrhizal (AM) fungi (Glomus mosseae and Glomus constrictum) and water stress on the growth and physiological performance of S. davidii seedlings under greenhouse pot conditions. Two soil water availability treatments (well-watered (WW) ?0.10 MPa; water-stressed (WS) ?0.86 MPa) were applied for 61 days. At the end of this experiment, G. mosseae and G. constrictum had colonized the roots of S. davidii seedlings. Water stress inhibited AM colonization, plant growth, chlorophyll concentration, gas exchange and chlorophyll fluorescence of S. davidii seedlings. Mycorrhizal seedlings had greater shoot dry weight, root dry weight, plant height, root length, instantaneous water use efficiency (iWUE), net photosynthetic rate (Pn), stomatal conductance (g_s), maximal photochemical efficiency of PSII photochemistry (Fv/Fm), lower intercellular CO₂ concentration and photochemical quenching values (qP), when compared with non-mycorrhizal seedlings under both WW and WS conditions. Furthermore, G. constrictum was found to be more efficient at improving the shoot and root mass, plant height, iWUE, Pn, g_s, qP, and ΦPSII of S. davidii seedlings, when compared with G. mosseae under both WW and WS conditions. Our results demonstrate that AM Glomus symbiosis enhanced S. davidii seedling resistance by improving its growth and physiological performance under water stress conditions. This suggests that Glomus inoculation is a potential tool for enhancing outplanting performance of S. davidii in semiarid areas of China.     

Effects of the physical properties of sphagnum peat on water stress in containerized picea mariana seedlings under simulated field conditions

Containerized black spruce seedlings, grown in Sphagnum peat at two degrees of humification (light and moderate) and two artificially created textural grades (with or without particles <1.3 mm), were subjected to controlled degrees of water stress using polyethylene glycol (PEG) at 0, 20, 75 and 175 g PEG kg^?1 H₂O, with and without dialysis membranes. After a 4‐day exposure, shoot water potential and water potential at the soil‐root interface were strongly affected by the humification of the peat at the highest PEG concentration. Differences in fine particle content of the peats had little effect on the physiological variables measured. Needle conductance was affected greatly by the presence of the dialysis membrane itself. The results suggest that high aeration, one of the most desirable physical properties of a growing medium for nursery production, may become undesirable once the seedlings are outplanted.     

Diplodia shoot blight and asymptomatic persistence of Diplodia pinea on or in stems of jack pine nursery seedlings

Diplodia pinea (syn. Sphaeropsis sapinea) is known as a major cause of damage to red pine (Pinus resinosa) seedlings in nurseries. The fungus can also be a latent pathogen of red pine seedlings, persisting in the absence of gross symptoms and later proliferating under conditions that induce host stress. In the fall of 2004, three nurseries in Wisconsin were surveyed to determine the potential for the occurrence of Diplodia shoot blight on jack pine (Pinus banksiana) seedlings and the persistence of D. pinea on or in asymptomatic seedlings of this species. Incidence of shoot blight was quantified in five 1 m long segments of an interior row in each of two survey areas in each nursery. The pathogen was identified on symptomatic seedlings collected in these areas on the basis of presence of characteristic pycnidia and conidia. Five groups of 20 asymptomatic seedlings were also collected in each of the two survey areas in each nursery. A segment of the lower stem/root collar from each of these asymptomatic seedlings was surface‐disinfested and culturally assayed using tannic acid agar. The mean incidence of shoot blight (as high as 9%) and mean frequency of cultural detection from asymptomatic seedlings (as high as 20%) were greatest in proximity to red pine windbreaks which are a source of inoculum. Only D. pinea was confirmed from subsets of symptomatic and asymptomatic seedlings which were tested using mt SSU rDNA polymerase chain reaction (PCR) primers that allow differentiation of D. pinea from D. scrobiculata and other fungi in the genus Botryosphaeria and related anamorphic fungi. Jack pine seedlings inoculated with D. pinea isolates obtained from asymptomatic nursery seedlings developed shoot blight symptoms in greenhouse trials. Thus, the ability of D. pinea to damage jack pine seedlings in nurseries has been documented and the potential for virulent strains of this latent pathogen to be distributed on asymptomatic jack pine seedlings from nurseries has been confirmed.