Estimating seed quantity and quality in shortleaf pine (Pinus echinata Mill.) cones from natural stands

The quantity and quality of seeds were studied in 886 shortleaf pine cones collected from 48 trees in each of two 15-ha natural stands in western Arkansas, USA. The number of seeds per cone ranged from 0 to 102, and averaged 34.6 and 38.0 in the two stands. Between 30 and 50 percent of the seeds were sound. Overall mean values for total number and total sound seeds compare with previous work. However, the wide variation between individual trees in a stand needs to be considered when evaluating cone abundance and estimating potential seed yields. For the stands observed in this study, a sample of 65 to 85 cones would be needed to estimate the number of sound seeds per cone (within ±3 seeds) with 95 percent probability. Sampling 1 or 2 cones from each of about 3 trees per ha would provide a reliable estimate of sound seed contents.     

The effects of acid rain and ozone on biomass and leaf area parameters of shortleaf pine (Pinus echinata Mill.)

Shortleaf pine (Pinus echinata Mill.) seedlings in 24 open-top chambers were exposed to combinations of ozone (carbon-filtered (control), ambient, 1.7 x ambient, and 2.5 x ambient) and acidic precipitation (pH 5.3, 4.3 and 3.3) for 16 months (1989 harvest) or 28 months (1990 harvest). Although the effects of acid rain were generally not significant, there was a trend toward increased aboveground biomass and leaf area in seedlings subjected to the low pH treatments. Because N concentrations in the soils generally increased with decreasing pH, we concluded that the effects of acid rain on aboveground biomass and leaf area were a consequence of an increasing concentration of soil N. In the 1989 harvest, seedlings in the 2.5 x ambient ozone treatment had significantly less biomass in all aboveground plant components and significantly less total leaf area than seedlings in the 1.7 x ambient ozone treatment. In the 1990 harvest, there were no significant effects of ozone on total aboveground biomass, although there was a trend toward reduced biomass in seedlings in the 2.5 x ambient ozone treatment. Both total leaf area and leaf biomass were significantly less in seedlings exposed to 2.5 x ambient ozone for 28 months than in both control seedlings and seedlings in the 1.7 x ambient ozone treatment. The greater, but not always significant, aboveground biomass and leaf area of seedlings in the 1.7 x ambient ozone treatment compared with control seedlings may be associated with the observed increase in soil nitrate concentration as a result of increased rates of leaf senescence and litterfall.     

Vascular cambial sucrose metabolism and growth in loblolly pine (Pinus taeda L.) in relation to transplanting stress

Sucrose synthase (SS) was the dominant enzyme of sucrose metabolism in both stem and root vascular cambial zone tissues of nursery-grown loblolly pine (Pinus taeda L.) seedlings. Acid invertase (AI) and neutral invertase (NT) activities were generally less than 10% of the SS activity in both tissues. In both cambial tissues, seasonal patterns of enzyme activity were observed for SS but not for AI or NI. The seasonal patterns of SS activity in stem and root cambia paralleled the periodic growth of stems and roots. Stems had high SS activity and growth during summer and early fall. Roots had substantial SS activity and growth during summer and fall, but SS activity and growth were even higher in winter. When seedlings were transplanted, about eight months elapsed before stem and root cambia resumed rates of growth and sucrose metabolism similar to those in control nontransplanted seedlings. Two months after transplanting, root SS was at its lowest, whereas AI activity in transplants was 50% higher than in control nontransplanted seedlings. In stems, SS activity decreased in response to transplanting, whereas AI and NI activities did not change appreciably. In loblolly pine tissues, SS was specific for uridylates, whereas the nucleotide triphosphate-dependent phosphofructokinase (NTP-PFK) had similar activity with either UTP or ATP. Except in winter, the NTP-PFK was less active than the pyrophosphate-dependent phosphofructokinase (PPi-PFK) during all seasons. The PPi-dependent PFK activity in nontransplanted seedlings followed similar seasonal and spatial patterns to those of SS activity. In actively growing tissues, such as stem cambial tissues in summer and root cambial tissues in winter, the measured total PFK to SS ratio ranged between 1.5/1 and 3/1. In contrast, in less actively growing tissues or transplanted seedlings, a greater decrease occurred in SS than in PFK activity, hence the ratio rose to as high as 12/1. It was concluded that: (1) SS was the dominant enzyme for sucrose metabolism in root and stem cambial tissues of loblolly pine seedlings; (2) both SS and PPi-PFK in the cambial tissues can be used as biochemical indicators of growth sink strength in stems and roots; and (3) both enzymes can be used as indicators of seedling stress caused by events such as transplanting and winter freezing.     

Improved recruitment and early growth of Scots pine (Pinus sylvestris L.) seedlings after fire and soil scarification

The success of seedling recruitment of Scots pine (Pinus sylvestris L.) is strongly dependent on soil surface properties, such as humus depth and moisture content. In an undisturbed forest floor, seedlings are seldom able to become established due to the high incidence of desiccation in the organic soil layer. Methods that remove the organic soil layer are often necessary to improve the availability for radicles to reach the more stable moisture regime in the mineral soil. In this study we investigated pine-seedling establishment after mechanical soil scarification, burning of litter (O_L) and burning of litter and humus (O_L and O_FH) in two mature pine stands in Germany. The herbaceous layer of the first stand was dominated by grasses (Molinia caerulea L. and Deschampsia flexuosa L.), whereas the herbaceous layer of the second stand was dominated by blueberry (Vaccinium myrtillus L.). Pine seeds were placed in experimental plots, and seedling numbers and heights were recorded at regular intervals. All treatments that removed organic soil resulted in higher seedling counts than did the undisturbed forest floor. The highest seedling counts were found on scarified and severely burnt plots, whereas seedling counts were lower on lightly burnt plots. Seedlings were significantly taller on burnt plots. This study shows that pine regeneration is stimulated by fire, not only in boreal forests, but also under central European conditions. With the expectation of higher fire frequency in the near future due to climatic changes, natural regeneration and succession on burnt sites should receive more focus in forest management and research.     

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.     

Maturation of maritime pine (Pinus pinaster Ait.) seedlings after exposure to a period of continuous light

Nine half-sib families of maritime pine (Pinus pinaster Ait.) of known adult performance were grown in continuous light at either 25 degrees C or 25/20 degrees C for 18 weeks. They were then exposed to a dormancy induction period followed by a dormancy release period and then grown for a further 9 weeks in a 16-h photoperiod at a day/night temperature of 25/20 degrees C. Seedlings exhibited great diversity in morphology at the end of the first growth period. The number of morphogenetic cycles varied between one and three and the form of the apical meristem ranged from a typical rosette to an adult-like bud. The type of seedling obtained at the end of the first growth period strongly influenced later growth, independently of the temperature regime. Maturity was proportional to the number of morphogenetic cycles achieved during the first growth period and was characterized by short growth duration, small primary needles and a high degree of fixed growth. The state of the apical meristem that underwent the dormancy period had less influence on the rate of maturation than the number of morphogenetic cycles. The time course of maturation was endogenously controlled and varied among traits. Conspicuous morphological differences were not associated with changes in the relationship between growth components at the phenotypic level. However, there seemed to be a shift in the genetic correlations between growth components after first budset.     

Research versus operational correlations between seedling survival and root growth potential of shortleaf pine

Relationships between root growth potential (RGP) and plantation survival were evaluated for 12 families of shortleaf pine (Pinus echinata Mill.) from Oklahoma and Arkansas in 2 consecutive years. Bare-root, 1+0 seedlings were either planted one day after lifting (nonstored) or were stored at 1° to 3°C for 4 weeks. Correlations were obtained using the traditional research method where seedlings are planted in the field prior to obtaining results from the RGP test. In addition, operational correlations were derived by planting seedlings after obtaining RGP test data.For nonstored seedlings, research correlations were generally higher (r values were larger in 18 of 24 cases) than operational correlations. The predictive ability of operational correlations may not be high enough with 28-day pot tests to warrant batch culling of freshly lifted shortleaf pine. However, results were mixed for stored seedlings. In one year, operational correlations were generally higher than research correlations.     

Regrowth and dry matter allocation in Quercus robur (L.) seedlings root pruned prior to transplanting

Two-year seedlings of Quercus robur (pedunculateoak) were root-pruned before transplanting in order toevaluate the importance of coarse versus fine roots onregrowth. Root systems were pruned by leaving c. 19,13 or 7 cm root from the root collar. Alternatively,coarse roots (>2 mm in diameter) were removed, leaving only the taproot and the fine roots (<2 mm)attached, or fine roots were removed from coarse rootsand taproot. Growth of shoots and roots after onegrowing season was compared to an unpruned controlunder standard nursery conditions. Seedlings rootpruned to 19, 13 or 7 cm were further tested undercompetition achieved by transplanting into a mixtureof clover and grass. Pruning of the root systemsignificantly reduced regrowth in terms of total plantDW in accordance with the severity of pruning, shootDW being more affected than root DW. Removal of coarseroots depressed final root DW whereas removal of fineroots reduced shoot DW and hence root:shoot ratioincreased. The study suggests that fine and coarseroots have different roles in root:shoot allocation.The competition test increased root:shoot ratioindicating that competition induced seedlings toallocate more of their resources into growth of theroot system.     

Carbon allocation, root exudation and mycorrhizal colonization of Pinus echinata seedlings grown under CO(2) enrichment

Increased exudation of carbon compounds from roots may provide a mechanism for enhancement of nutrient availability to plants growing in a CO(2)-enriched atmosphere. Therefore, the effect of atmospheric CO(2) concentration on carbon allocation and root exudation was investigated in Pinus echinata Mill. (shortleaf pine) seedlings. After 34 and 41 weeks, seedlings growing in 695 microl l(-1) CO(2) allocated proportionately more (14)C-labeled photosynthate to fine roots than did seedlings growing in ambient air. This was associated with greater fine root mass and mycorrhizal density in CO(2)-enriched plants after 34 weeks. Exudation of soluble, (14)C-labeled compounds from roots also was greater in these plants at 34 weeks, but the effect of CO(2) concentration on exudation did not persist at 41 weeks.     

Short-day treatment affects growth,physiological parameters and needle proteome of Chinese pine (Pinus tabuliformis Carr.) seedlings

New Forests - Previous studies have proven that short-day treatment is an effective means to regulate shoot growth and abiotic stress resistance in conifers. However, the effects on Chinese pine...     

Effects of nitrogen fertilization and temperature on frost hardiness of Aleppo pine (Pinus halepensis Mill.) seedlings assessed by chlorophyll fluorescence

In a 14-week study, 1-year-old Aleppo pine seedlings were grownin two growth chambers. Seedlings were artificially hardenedby decreasing photoperiod and temperature. In each chamber halfof the seedlings were fertilized with nitrogen (8.4 mg seedling^–1).In order to determine the relative importance of the hardeningenvironment versus fertilization, each chamber was programmedto decrease night temperatures down to a low of 8 or 4°C.Chlorophyll fluorescence and frost hardiness was measured fivetimes during the experiment. A sample of seedlings from eachtreatment was exposed to an artificial frost at –5°Cand the freezing effects were assessed by measurements of chlorophyllfluorescence and visual evaluation of needle damage. Seedlingsincreased their frost hardiness during the experiment in allthe treatments but the ratio of variable to maximal chlorophyllfluorescence (F_v/F_m) measured before freezing did not vary duringthe experiment. This indicates that Aleppo pine maintains itsphotosynthetic ability during hardening in contrast to otherconiferous species from colder climates. The effect of nitrogenfertilization on frost hardiness was small in comparison withchamber effect. Nitrogen fertilization slightly delayed theacquisition of hardening in the coldest chamber. Seedlings inthe warmest chamber did not become fully resistant to –5°C,but in the coldest chamber, where night temperature reached4°C, all the seedlings were resistant to the frost. Severedamage caused by frost could be related to a rapid rise of minimalfluorescence (F₀) but the best index of damage was the dropof F_v/F_m after freezing.     

Growth and visible responses of Scots pine (Pinus sylvestris L.) seedlings to simulated summer frost

Scots pine seedlings, 20–30 days old, were exposed to simulated summer frost in controlled environment growth chambers. The responses observed showed extreme variation between individual seedlings; seedlings suffering from needle-necrosis, healthy-looking, and dead seedlings were found in the same treatment. Four days exposure to temperatures below –4.5°C caused multiple-leaders maximally in 7.7 % of the pine seedlings and resulted in decreased shoot and root dry weight and shoot length.     

Exposure to a pine pathogen enhances growth and disease resistance in Pinus radiata seedlings

Most studies of Fusarium circinatum, the cause of pitch canker in pines, have focused on its activity as a pathogen. However, recent findings indicate that this fungus can colonize roots of Pinus radiata without inducing symptoms. Contrary to expectations, this study revealed that seedlings grown in infested sand grew more rapidly than seedlings not exposed to F. circinatum, based on root and shoot biomass, with modifications to root system architecture, including increased mycorrhizal root development. These effects were dependent on inoculum density and duration following growth in infested rooting medium. Plants exposed to F. circinatum expressed elevated resistance to stem infections, which significantly decreased the incidence of mortality; as above, effects were dependent on inoculum density. Resistance to stem infections was also enhanced in seedlings that emerged through infested litter, as occurs in native stands. Beneficial to neutral interactions of F. circinatum with its host suggest that the life history of this fungus may be more complex than previously recognized, with activities similar to non‐pathogenic endophytes. The potential for non‐lethal infections by F. circinatum to induce resistance in seedlings may influence dynamics of stand establishment. Overall, these results indicate that pathogenic organisms with asymptomatic states may have cryptic ecological functions that extend beyond the impacts of disease.     

Photosynthetic acclimation of shade-grown red pine (Pinus resinosa Ait.) seedlings to a high light environment

Photosynthetic light acclimation in red pine (Pinus resinosa Ait.) seedlings was examined in a greenhouse study to better understand the physiological response of this species to increased light intensity following release from competition. Seedlings grown in a high (HL), medium (ML) or low (LL) light environment for 12 weeks were transferred to high light. Gas exchange and chlorophyll fluorescence of ML and LL seedlings were measured prior to and following transfer and compared with the HL control treatment. Photosynthetic characteristics were related to initial light treatment and time after transfer. Acclimation of gas exchange features to high light in shade formed ML and LL foliage was relatively rapid, with similar values among light treatments within 57 days of transfer. Acclimation of net photosynthetic rate was similar in ML and LL seedlings, and was associated primarily with increased mesophyll conductance to CO₂. The ratio of variable to maximal chlorophyll fluorescence (F_v/F_m) decreased initially after transfer, especially in LL seedlings, but recovered to normal values after 57 days. Red pine seedlings appear to be well adapted for photosynthetic acclimation to high light intensity, consistent with that reported for other early successional tree species.     

Effect of early transplanting upon growth and development of spruce and pine seedlings in paperpot containers

Black spruce (Picea mariana [Mill.] B.S.P.) and jack pine (Pinus banksiana Lamb.) seedlings were transplanted into Japanese paperpots at five different stages of development between the occurrence of an erect hypocotyl and 14 days after initiation of primary needles. Transplanting difficulty increased between the first and last treatments, and transplanting generally became impracticable after the stage of primary-needle initiation. Transplanting had a relatively minor effect on the eventual size of 12-week-old pine and 16-week-old spruce seedlings, although the incidence of root deformities inside the container increased rapidly as transplanting was delayed beyond the erect-hypocotyl stage. These deformities persisted after seedlings were outplanted, but did not appear to have a negative effect on the subsequent growth of new roots into the planting medium. Despite the absence of major adverse effects, it is recommended that transplanting be avoided as a routine practice for supplementing stocking in trays of containerized seedlings because of possible biological risks, the difficulty of ensuring careful transplanting and high labor costs.     

Isolation,characterization and effect of plant-growth-promoting rhizobacteria on pine seedlings(Pinus pseudostrobus Lindl.)

In this study,10 bacterial strains were isolated from the rhizosphere of coniferous trees on Mount Tla′loc in Mexico.The strains were characterized by their capacity to produce auxins,solubilize phosphates and stimulate mycelial growth of the ectomycorrhizal fungus Suillus sp.All isolates were identified at the molecular level.Moreover,an experiment was established to evaluate the response of Pinus pseudostrobus seedlings to inoculation with the rhizobacteria strains.The isolated strains belonged to the species Cupriavidus basilensis,Rhodococcus qingshengii,R.erythropolis,Pseudomonas spp.,P.gessardii,Stenotrophomonas rhizophila and Cohnella sp.All of the strains produced auxins;the best producer was R.erythropolis CPT9(76.4 lg mL^-1).P.gessardii CPT6 solubilized phosphate at a significant level(443 lg mL^-1).The strain S.rhizophila CPT8 significantly increased the radial growth of the ectomycorrhizal fungus Suillus sp.by 18.8%.Five strains increased the dry mass of the shoots;R.qingshengii CPT4 and R.erythropolis CPT9 increased growth the most,by more than 20%.Inoculation with plant-growth-promoting rhizobacteria can be a very useful practice in a forest nursery to produce healthy,vigorous plants.     

Acclimation of loblolly pine (Pinus taeda) seedlings to high temperatures

To determine the extent to which loblolly pine seedlings (Pinus taeda L.) acclimate to high temperatures, seedlings from three provenances-southeastern Texas (mean annual temperature 20.3 degrees C), southwestern Arkansas (mean annual temperature 16.2 degrees C) and Chesapeake, Maryland (mean annual temperature 12.8 degrees C)-were grown at constant temperatures of 25, 30, 35 or 40 degrees C in growth chambers. After two months, only 14% of the seedlings in the 40 degrees C treatment survived, so the treatment was dropped from the experiment. Provenance and family differences were not significant for most measured variables. Total biomass was similar in the 25 and 30 degrees C treatments, and less in the 35 degrees C treatment. Foliage biomass was higher, and root biomass lower, in the 30 degrees C treatment compared with the 25 degrees C treatment. Net photosynthesis and dark respiration of all seedlings were measured at 25, 30 and 35 degrees C. Both net photosynthesis and dark respiration exhibited acclimation to the temperature at which the seedlings were grown. For each temperature treatment, the highest rate of net photosynthesis was measured at the growth temperature. Dark respiration rates increased with increasing measurement temperature, but the basal rate of respiration, measured at 25 degrees C, decreased from 0.617 &mgr;mol m(-2) s(-1) in the 25 degrees C treatment to 0.348 &mgr;mol m(-2) s(-1) in the 35 degrees C treatment, resulting in less carbon loss in the higher temperature treatments than if the seedlings had not acclimated to the growth conditions. Temperature acclimation, particularly of dark respiration, may explain why total biomass of seedlings grown at 30 degrees C was similar to that of seedlings grown at 25 degrees C.     

Canopy fuel characteristics and potential crown fire behavior in Aleppo pine (Pinus halepensis Mill.) forests

Canopy fuel characteristics that influence the initiation and spread of crown fires were measured in representative Aleppo pine (Pinus halepensis Mill.) stands in Greece. Vertical distribution profiles of canopy fuel load, canopy base height and canopy bulk density are presented. Aleppo pine canopy fuels are characterized by low canopy base height (3.0–6.5 m), while available canopy fuel load (0.96–1.80 kg/m²) and canopy bulk density (0.09–0.22 kg/m³) values are similar to other conifers worldwide. Crown fire behavior (probability of crown fire initiation, crown fire type, rate of spread, fireline intensity and flame length) in Aleppo pine stands with various understory fuel types was simulated with the most updated crown fire models. The probability of crown fire initiation was high even under moderate burning conditions, mainly due to the low canopy base height and the heavy surface fuel load. Passive crown fires resulted mostly in uneven aged stands, while even aged stands gave high intensity active crown fires. Assessment of canopy fuel characteristics and potential crown fire behavior can be useful in fuel management and fire suppression planning.     

Movement of respiratory CO(2) in stems of loblolly pine (Pinus taeda L.) seedlings

Temperature-independent fluctuations in stem CO(2) efflux were measured in Pinus taeda L. seedlings. Stem CO(2) efflux was measured during high and low transpiration rates, high and low net photosynthesis rates, and normal and interrupted substrate supply conditions. Stem CO(2) efflux rates were an average of 6.7% lower during periods of high transpiration compared to periods of low transpiration. This difference in stem CO(2) efflux rates was not due to water stress. The most likely cause was movement of respiratory CO(2) in the transpiration stream. Interruption of substrate supply to the stem by phloem girdling reduced stem CO(2) efflux rates. Increasing net photosynthesis rates from low to high had no effect on stem CO(2) efflux, but decreasing net photosynthesis from high to low caused relatively small reductions in stem CO(2) efflux. These results indicate that diurnal changes in net photosynthesis rate may play a small role in temperature-independent afternoon depressions of stem CO(2) efflux. The transport of respiratory CO(2) by the transpiration stream compromises measurements of woody tissue respiration obtained by commonly accepted gas exchange techniques. This phenomenon could also affect measurement of leaf net photosynthesis and branch woody tissue respiration.     

Large-scale genetic structure and drought-induced effects on European Scots pine (Pinus sylvestris L.) seedlings

Seedlings of ten provenances of Scots pine (Pinus sylvestris L.) from the southwestern to the central part of the species distribution range were investigated in a greenhouse experiment under dry conditions compared to a well-watered control. We conducted an isozyme study and recorded phenology as well as growth traits during the first year of growth. Genetic variability and genetic diversity increased from the westernmost provenances to those central to the distribution. Provenances from the Apennines and Spain revealed the strongest genetic differentiation compared to all others, whereas populations from Germany, Poland and Bulgaria were found to belong to the same gene pool. Seedling development and bud set were faster in northern populations. Shoot length was highest for Polish and German provenances within both regimes, populations from France and Spain had the longest roots. Reduced soil moisture slowed later development stages and delayed bud set for all provenances by about 5 days. Shoot growth decreased considerably under the dry regime for all provenances. German provenances were the most sensitive to drought, whereas Spanish provenances showed adaptation to drought conditions indicated by the lowest reduction in shoot growth relative to optimum conditions. The results of this pilot study show that under more pronounced drought conditions with repeated drought events, the so far established superiority of northeastern provenances compared to southwestern ones could diminish in the future.