Responses of Pinus clausa, Pinus serotina and Pinus taeda seedlings to anaerobic solution culture. I. Changes in growth and root morphology

Seedlings of pond pine ( Pinus serotina Michx.), sand pine [ P. clausa (Engelm.) Sarg.] and two edaphic seed sources of loblolly pine ( P. taeda L., dry- and wet-site seed sources) were grown in non-circulating, continuously flowing solution culture under aerobic (250 μ M O₂) and anaerobic (≤ 23 μ M O₂) conditions. Survival was 100% for all seedlings at 11 weeks. Although shoot height, biomass and leaf emergence of loblolly and pond pine seedlings were not significantly affected by 15 or 30 days of anaerobic growth conditions, root biomass was significantly reduced. Sand pine suffered the largest reduction in biomass, showing extensive root dieback and shoot chlorosis with retarded leaf development. These anaerobically induced symptoms of flooding injury were less severe in dry-site loblolly pine, and absent in wet-site loblolly and pond pine seedlings. Adventitious or new, secondary roots, and stem or taproot lenticels were particularly abundant under the 30-day anaerobic treatment in wet-site loblolly and pond pine seedlings, present to a lesser degree in dry-site loblolly pine, and nearly absent in sand pine seedlings. These results indicate that much of the immediate damage from flooding is due to the anoxic condition of the root rather than to the build-up of phytotoxins or soil nutrient imbalances. On the basis of overall seedling vigor, root plasticity and growth, we suggest the following flooding-tolerance/intolerance species (seed source) gradient: pond pine ≥ wet-site loblolly pine > dry-site loblolly pine > sand pine.     

Aerenchyma and lenticel formation in pine seedlings: A possible avoidance mechanism to anaerobic growth conditions

Seedlings of pond pine ( Pinus serotina Michx.), sand pine [ P. clausa (Engelm.) Sarg.], and loblolly pine ( P. taeda L., wet-site and drought-hardy seed sources) were grown in hydroponic solution culture using a non-circulating, continuously flowing design under anaerobic or aerobic conditions to determine whether flooding tolerance was correlated with enhanced internal root aeration. Transport of atmospheric O₂ from the shoot to the root of anaerobically grown loblolly and pond pine seedlings was demonstrated via rhizosphere oxidation, using both reduced indigo-carmine solution and a polarographic, ensheathing Pt-electrode. Stem and root collar lenticels were the major sites of atmospheric O₂ entry for submerged roots in these seedlings. No O₂ leakage was detected from roots of aerobically grown pine seedlings. Longitudinal and radial pathways for gaseous diffusion via intercellular air spaces in the pericycle and between ray parenchyma cells, respectively, were demonstrated histo-logically in anaerobically grown loblolly and pond pines. Rhizosphere oxidation, and lenticel and aerenchyma development in roots of flood-intolerant sand pine seedlings grown in anaerobic solutions were minimal. Only 15 days of anaerobic growth conditions were necessary to increase internal root porosities of loblolly and pond pine seedlings – although not to the extent found in seedlings treated for 30 or 75 days. Histological results indicated that root tissue in the secondary stage of growth was capable of forming intercellular air spaces, demonstrating a degree of internal plasticity – at least in the more flood-tolerant loblolly and pond pine seedlings.     

Modelling Inter- and Intra-specific Competition in Loblolly Pine (Pinus taeda L.) Plantations on Cutover, Site-prepared Lands

The presence of volunteer hardwood species in loblolly pineplantations demands studies on both intra- and inter-specificcompetition in order to build growth and yield models and toguide vegetation management for these stands. This paper, basedon analyses of data collected from a thinning study, reportsan investigation of responses of loblolly pine and hardwoodspecies towards intra- and inter-specific competition. Underhigh levels of overall competition, hardwood species were morecompetitive, both intra- and inter-specifically. Intra-specificcompetition was more effective in reducing hardwood basal areagrowth than inter-specific competition under high levels ofoverall competition. However, under low levels of overall competition,intra- and inter-specific competition were quantitatively similartoward loblolly pine basal area growth. The basal area growthof hardwoods was significantly related to levels of inter-specificcompetition, but not with intra-specific effects under low levelsof overall competition. More than half of the variability inloblolly pine basal area growth under unthinned control andlight thin treatments could be accounted for by competitioneffects. However, only one third of the variability could beexplained for loblolly pine under heavy thin treatment. Forhardwood species, the percentage of growth variation accountedfor by competition was about 45%, and did not change among thethinning treatments. Different resource demands between thetwo categories of tree species and a certain amount of thinningshock were suggested by the modelling results.Copyright 1994,1999 Academic Press Neighbourhood approach, competition index, hardwood, no-interaction model, basal area, reciprocal yield law     

Responses of Pinus clausa, Pinus serotina and Pinus taeda seedlings to anaerobic solution culture. II. Changes in tissue nutrient concentration and net acquisition

Fifteen or 30 days of anaerobic growth conditions significantly reduced shoot and root nitrogen, potassium, phosphorus, iron and manganese concentrations in seedlings of pond pine ( Pinus serotina Michx.), sand pine [ P. clausa (Engelm.) Sarg.] and drought-hardy and wet-site loblolly pine ( P. taeda L.) grown in a culture system using non-circulating, continuously flowing solution. Calcium and shoot magnesium levels were least affected by anaerobic growth conditions – largely reflecting the passive nature of their uptake. Shoot and root nutrient content (mg nutrient pot^-1) followed similar trends, with wet-site loblolly and pond pine seedlings least affected by anaerobic solution culture. Shoot biomass of wet-site loblolly and pond pine seedlings was not affected by anaerobiosis, suggesting an increase in shoot nutrient utilization efficiency. Root biomass was significantly reduced by 15 or 30 days of anaerobiosis, with sand pine exhibiting the largest reduction in root dry weight (57%).
These results suggest that anaerobiosis interferes with net nutrient acquisition, even under the high nutrient conditions provided by solution culture. Sand pine suffered the largest reductions in shoot and root biomass and nutrient concentrations, showing earlier symptoms of waterlogging injury and nutrient stress than drought-hardy loblolly pine seedlings. Whether net nutrient acquisition decreased because of the reduction in root surface area available for absorption and/or reduced uptake efficiency cannot be ascertained from these data.     

Longleaf and loblolly pine seedlings respond differently to soil compaction,water content,and fertilization

Aims

Longleaf pine (Pinus palustris Mill.) is being restored across the U.S. South for a multitude of ecological and economic reasons, but our understanding of longleaf pine’s response to soil physical conditions is poor. On the contrary, our understanding of loblolly pine (Pinus taeda L.) root and shoot growth response to soil conditions is well established.

Methods

We performed a comparative greenhouse study which modeled root length density, total seedling biomass, and the ratio of aboveground:belowground mass as functions of volumetric water content, bulk density and soil fertility (fertilized or not).

Results

Root length density was about 35 % greater in longleaf pine seedlings compared to loblolly pine seedlings, and was reasonably well modeled (R ²?=?0.54) for longleaf pine by bulk density (linear), volumetric water content (quadratic), soil fertility, and the interactions of bulk density, volumetric water content, species, and soil fertility. The aboveground:belowground mass ratio (ABR) increased at both extremes of water content.

Conclusions

This research indicates that young longleaf pine seedling root systems respond more negatively to extremes of soil physical conditions than loblolly pine, and compacted or dry loamy soils should be ameliorated in addition to normal competition control, especially on soils degraded by past management.     

SEASONAL TRENDS IN RATES OF PHOTOSYNTHESIS AND RESPIRATION OF LOBLOLLY PINE AND WHITE PINE SEEDLINGS

Mc Gregor , William H. Davis (Clemson Coll., Clemson, S.C.), and Paul J. Kramer . Seasonal trends in rates of photosynthesis and respiration of loblolly pine and white pine seedlings. Amer. Jour. Bot. 50(8): 760–765. Illus. 1963.—Seasonal trends in the rates of photosynthesis and respiration of potted loblolly and white pine seedlings were studied by measuring CO₂ exchange with an infra-red gas analyzer at intervals during a year. The seedlings were kept out-of-doors, but measurements were made indoors at 25 C and 4000 ft-c of light. Beginning in February, the rate of photosynthesis per plant of both species increased, reaching a peak in mid-September for loblolly pine and in mid-July for white pine and then declining rapidly after mid-September for both species. The spring increase began before new needles emerged and the autumn decline was not accompanied by appreciable loss of foliage. The respiration rate of loblolly pine increased steadily throughout the year. Respiration of white pine increased until mid-May and then remained fairly constant through the remainder of the year. Maximum rate of photosynthesis per unit of fascicle length occurred in July for the white pine and in May for the loblolly pine, and rates declined after September in both species. Respiration rate per unit of fascicle length showed a marked increase as spring growth began in April, then decreased to a minimum in September and increased during the winter in both species. Total chlorophyll per seedling reached a maximum in September for both species and declined slightly during winter. Total chlorophyll per unit of fascicle length increased in the spring, declined slightly in midseason, and increased again in September. Photosynthesis per unit of chlorophyll reached a maximum in May for loblolly pine and in July for white pine. Stem elongation of white pine was 88% completed by May 15. On the same date, stem elongation of loblolly pine was only 42% completed.     

Characterization of phosphorus uptake in slow- and fast-growing southern pine seedlings grown in solution culture

The effects of low-P growth conditions on growth and net P acquisition were examined in two species of pine that are indigenous to P-deficient soils of the Atlantic Coastal Plain: pond pine (Pinus serotina Michx.), a moderately-fast growing pine, and a slow-growing seed source of loblolly pine P. taeda L.) from Texas. Short-term ³²P uptake experiments were conducted using intact nonmycorrhizal seedlings that had been grown for 7 weeks in continuously-flowing solution culture at 5 or 100 µM P. Growth and P uptake of pond pine were more responsive to a higher P supply than the slow-growing loblolly pine. Pond pine seedlings in the 100 µM P treatment were twice the size of those grown in 5 µM P and accumulated almost five times as much seedling P. In contrast, seedling biomass of loblolly pine increased by only 8% under high-P growth conditions, and seedlings accumulated twice as much P, reflecting the higher P concentrations in shoot and root tissues. Although rates of unidirectional influx of ³²P were 22 and 61% higher under low-P growth conditions in pond and loblolly pine, respectively, net uptake rates in seedlings from the 5 µM P treatment were over three times those of seedlings grown in 100 µM P. These results suggest that unidirectional efflux out of the root was controlling net uptake of P as much, if not more, than unidirectional influx. Efflux of³² P out of root tissue, particularly older tissue, decreased in seedlings grown under low-P conditions, possibly due to a reduction in the size of the phosphorus pool available for efflux, i.e. the soluble P_i pool. Over 75% of the total root P in both loblolly and pond pine seedlings grown in 100 µM P treatment was present as organic P, suggesting that organic P, particularly phytate, may represent important storage pools in roots of woody species. Within each species, higher rates of influx and net uptake in seedlings from the low-P treatment were associated with lower P concentrations in shoot and root tissue, and shoot FW:root FW ratios. Efflux may represent a short-term means of regulating net P uptake, while the demand for P created by growth and storage may represent a long-term regulation.     

Morphology and tissue quality of seedling root systems of Pinus taeda and Pinus ponderosa as affected by varying CO2, temperature,and nitrogen

Rising atmospheric carbon dioxide, nitrogen deposition and warmer temperatures may alter the quantity and quality of plant-derived organic matter available to soil biota, potentially altering rates of belowground herbivory and decomposition. Our objective was to simulate future growth conditions for an early successional (loblolly) and late successional (ponderosa) species of pine to determine if the physical and chemical properties of the root systems would change. Seedlings were grown for 160 days in greenhouses at the Duke University Phytotron at 35 or 70 Pa CO₂ partial pressure, ambient or ambient + 5 °C temperature, and 1 or 5 mMNH₄O₃. Roots from harvested seedlings were analyzed for changes in surface area, specific root length, mass, total nonstructural carbohydrates (TNC), and concentrations of macro-nutrients. Surface area increased in both species under elevated CO₂, due primarily to increases in root length, and this response was greatest (+138%) in loblolly pine at high temperature. Specific root length decreased in loblolly pine at elevated CO₂ but increases in mass more than compensated for this, resulting in net increases in total length. TNC was unaffected and nutrient concentrations decreased only slightly at elevated CO₂, possibly from anatomical changes to the root tissues. We conclude that future growth conditions will enhance soil exploration by some species of pine, but root carbohydrate levels and nutrient concentrations will not be greatly affected, leaving rates of root herbivory and decomposition unaltered.     

Water relations of pine seedlings in relation to root and shoot growth

The effects of water stress on growth and water relations of loblolly and white pine seedlings were studied during series of drying cycles. As mean soil water potential decreased, growth of roots, needles, and buds decreased. Growth of roots during successive severe drying cycles was not uniform, however. A study of needle and root extension showed that of the total growth of roots for 3 7-day drying cycles, only 6% occurred during the third cycle, while needle extension was uniform for the 3 cycles. The difference in response of needles and roots to drying cycles may be attributed primarily to the effect of water stress on the growing region. When subjected to a severe stress, roots matured toward the tip and became dormant, resulting in less growth during subsequent drying cycles. The intercalary growing region of needles, however, was not altered seriously enough by the stress to cause a difference in amount of growth during each drying cycle.

Transpiration of loblolly pine was lower in the second drying cycle than in the first. Needle water potential after rewatering was as high as that of control plants watered daily; root resistance was apparently not important in restricting transpiration during a second drying cycle. Needle diffusion resistance of loblolly pine, measured with a low-resistance diffusion porometer, was slightly higher during the second drying cycle than during the first. In addition, many primary needles were killed during the first period of stress. These factors contributed to the reduction of transpiration during the second drying cycle. Diffusion resistance of Coleus increased and transpiration ceased during the first drying cycle while water potential remained relatively high. After rewatering, both leaf resistance and transpiration returned to the control level, presumably because the stress during the first period of drying was not severe. The diffusion resistances observed for well-watered plants were 30 to 50 sec·cm⁻¹ for loblolly pine, 3 to 5 sec·cm⁻¹ for Coleus, and 4 to 6 sec·cm⁻¹ for tomato. These values agree closely with those reported by other workers.

     

A Technique for Determining Quantitative Expressions of Dormancy in Seeds

A method of quantifying the heterogeneity of dormancy withinseed populations is described. The method compares germinationresponses of unstratified and stratified seeds of Pinus taedaL. (loblolly pine). Mathematical functions were fitted to cumulativegermination curves of unstratified and stratified seeds andfunctions were integrated to determine areas beneath respectivecurves. The area between respective curves was calculated bydifference and was considered to be a measure of dormancy withinseed lots. The method may be used to help identify various controlsof seed dormancy. seed germination, stratification, germination curves, Pinus taeda, loblolly pine, dormancy     

Micropropagation of shortleaf,Virginia and loblolly x shortleaf pine hybrids via organogenesis

Protocols were developed for the micropropagation of shortleaf pine (Pinus echinata Mill.), loblolly (P. taeda L.) x shortleaf pine hybrids, and Virginia pine (P. virginiana Mill.). For meristematic tissue induction, modified Gresshoff & Doy (GD) medium with a high concentration of benzyladenine (BA) and short pulse treatment was best for loblolly x shortleaf hybrids whereas a lower concentration of BA and longer pulse treatment was best for shortleaf and Virginia pines. Shoot growth rate for all species was generally slower on Schenk & Hildebrandt medium than on GD medium. Addition of activated charcoal improved shoot growth of shortleaf pine but not of Virginia pine or the loblolly x shortleaf hybrids. Separation of shoots was beneficial before placing in the advanced growth medium. Both GD and Litvay's media produced good advanced shoot growth, especially following the addition of 0.5% activated charcoal. Individual shoot heights of 2–3 cm and 8–12 weeks of age after separation from the cluster were best for rooting. Root induction declined rapidly thereafter. Modified GD medium with 0.5 mg 1^-1 -naphthaleneacetic acid plus 1.0 mg 1^-1 3-indolebutyric acid and 20 g 1^-1 sucrose was best for root induction for all species except shortleaf pine. Addition of activated charcoal produced better root systems. Too high a light intensity resulted in a lowered frequency of rooting. A large number of plantlets was produced.     

Adventitious root formation in cuttings of loblolly pine (Pinus taeda L .): developmental sequence and effects of maturation

 Adventitious root formation in cuttings from fascicular shoots in loblolly pine (Pinus taeda L.) consists of four more or less discontinuous stages: (1) proliferation of cells at the base of the cutting, (2) differentiation of wound vascular tissue and periderm, (3) dedifferentiation of a zone near the wound cambium and wound phloem to form a root initial, and (4) formation of a root meristem. Anatomical changes during adventitious root initiation are described in cuttings from donors of different types and ages. Cuttings from seedlings and 3- to 7-year-old hedged stock plants rooted better than cuttings from 3-year-old tree form donors. It is concluded that the loss of rooting capacity in loblolly pine can be arrested by shearing loblolly pine stock plants to low hedges. The process of root initiation, however, was similar in cuttings from all sources and is apparently not the cause for the rapid decline of rooting potential with increasing age of the donor plant. Received: 3 June 1997 / Accepted: 15 August 1997     

The Influence of Pine on the Form of Sitka Spruce Fine Roots

The form of fine roots of Sitka spruce (Picea sitchensis Bong.Carr.) when grown in immediate proximity to roots of Scots pine(Pinus sylvestris L.) and when grown separately was comparedusing split-root systems. When spruce roots were intimatelymixed with pine roots the mean length of individual spruce lateralswas significantly greater, while the length: weight ratio andnumber of root tips: weight ratio were smaller than when grownalone. A similar alteration in growth strategy was achievedby direct addition of mineral nitrogen. Key words: Fine root form, pine-spruce interactions     

An Individual Stand Growth Model for Mean Plant Size Based on the Rule of Self-thinning

A growth model for pure, even-aged stands of plants is asymptoticallybounded above by the self-thinning rule that relates maximumplant size to stand density. The model characterizes accretionin mean size as a deviation from the limiting size. It consistsof a function relating mean size to time and density and a companionsurvival model. The growth model is obtained by substitutingthe survival model for density in the mean size relationship.Model flexibility is demonstrated by fitting it to annual remeasurementsof mean size and number of plants per unit area in a stand ofPinus taeda L. 3/2-power rule, mortality, survival, stand dynamics, plant growth model, loblolly pine     

Average effect of a mutation in lignin biosynthesis in loblolly pine

Cinnamyl alcohol dehydrogenase (CAD, E.C. 1.1.1.195) is a monolignol biosynthetic enzyme that catalyzes the final step of lignin subunit biosynthesis in higher plants. Recently, a mutant allele of the cad gene, cad-n1, encoding for the CAD enzyme, was discovered in loblolly pine. By reducing the expression of the cad gene, this mutant has a decreased lignin content and major changes in the lignin composition in wood. In this study, we found that the substitution of a wild-type allele by cad-n1 was associated with a significant effect on 2nd-year shoot elongation in a half-sib family of loblolly pine (designated family 7–1037). The average effect of cad-n1 appeared to increase with tree growth and was greater for stem radial growth than height growth. An increase of 14.1% in de-barked volume in year 4 was associated with cad-n1. Co-segregation analysis indicated that the cad locus itself might represent a gene that governs stem growth in pine. The significance of the mutation cad-n1 for tree growth and wood processing is discussed. Received: 31 December 1998 / Accepted: 30 January 1999     

Shoot--root Plasticity and Episodic Growth in Red Pine Seedlings

Red pine seedlings of a half-sib seed source were grown in growthchambers under thermoperiodic regimes of 30/20 °C, 25/15°C and 20/10 °C day/night temperatures. Classical growthanalyses based on weekly harvests of leaves, stem and rootswere employed to study the first 3 to 15 weeks of seedling development.Leaf and root growth were inversely related and episodic. Significantshort term surges in growth of either organ were effective inreversing periodic imbalances that occurred, thus maintaininga long term dry weight equilibrium between above and below groundseedling parts. Adaptive plasticity in the leaf-root balanceat different temperatures gave plants grown at 25/15 °Ca larger proportion of leaves relative to roots and a greatersize compared to seedlings grown under other regimes. Episodicfluctuations in leaf and root growth occurred simultaneouslywith depressions in net assimilation rate. Apparently, balancedgrowth is maintained at an assimilatory cost to the plant, periodic‘corrections’ of shoot—root imbalances requiringcarbohydrate conversion and energy expenditure. Pinus resinosa Ait., red pine, episodic growth, shoot—root balance, plasticity, net assimilation rate, growth analysis     

Regulation of loblolly pine (Pinus taeda L.) arginase in developing seedling tissue during germination and post-germinative growth

After seed germination, hydrolysis of storage proteins provides a nitrogen source for the developing seedling. In conifers the majority of these reserves are located in the living haploid megagametophyte tissue. In the developing loblolly pine (Pinus taeda L.) seedling an influx of free amino acids from the megagametophyte accompanies germination and early seedling growth. The major component of this amino acid pool is arginine, which is transported rapidly and efficiently to the seedling without prior conversion. This arginine accounts for nearly half of the total nitrogen entering the cotyledons and is likely a defining factor in early seedling nitrogen metabolism. In the seedling, the enzyme arginase is responsible for liberating nitrogen, in the form of ornithine and urea, from free arginine supplied by the megagametophyte. In this report we investigate how the seedling uses arginase to cope with the large arginine influx. As part of this work we have cloned an arginase cDNA from a loblolly pine expression library. Analysis of enzyme activity data, accumulation of arginase protein and mRNA abundance indicates that increased arginase activity after seed germination is due to de novo synthesis of the enzyme. Our results suggest that arginase is primarily regulated at the RNA level during loblolly pine seed germination and post-germinative growth.