Nursery Culture Impacts Cold Hardiness in Longleaf Pine (Pinus palustris) Seedlings

Success in restoring longleaf pine ecosystems depends on outplanting high‐quality longleaf pine seedlings. One important and relatively understudied attribute of seedling quality is cold hardiness. A suite of trials was conducted to investigate the influence of common nursery cultural practices on longleaf pine cold hardiness. Cold hardiness was increased with higher rates of nitrogen, unaffected by copper coating containers, greater for foliage than for root‐collar tissue, and tended to increase with increases in container size.     

Localization and Activity of Naphthylamidases in Germinating Seeds of Scots Pine, Pinus sylvestris

Extracts prepared from endosperms of germinating seeds of Scots pine, Pinus sylvestris L., rapidly hydrolysed the β-naphthylamides of L-phenylalanine and L-leucine optimally at pH 6.5 and that of L-arginine at pH 7.7. Disc electrophoresis followed by activity staining showed that the activities were due to two naphthylamidases (aminopeptidases) with different substrate specificities. Seeds were allowed to germinate at 20°C on agar gel in the dark and the activities on the three substrates were assayed from separated endosperms and seedlings at various stages of germination. The activities in the endosperm of resting seeds were relatively high and they remained unchanged throughout the period of reserve protein mobilization (seedling length up to 50 mm), after which they began to decrease. The activities of the naphthylamidases are rather small compared with those of the two alkaline peptidases of pine, contributing about 17% of the total amino-peptidase activity in the endosperm of germinating seeds. The total aminopeptidase activity is sufficient to account for the rate of storage protein mobilization during germination. In the seedlings the naphthylamidase activities (per seedling) increased continuously during germination, and activities per g dry weight were higher than those in the endosperm.     

Characterization of the Proteinases Present in Germinating Seeds of Scots Pine, Pinus sylvestris

Methods were developed to determine proteinase activity in germinating seeds of Scots pine. The assays were based on the liberation of TCA-soluble peptides from haemoglobin at pH 3.7 and from casein at pH 5.4 and pH 7.0; the reaction products were determined by the Lowry method. — Endosperms separated from seeds at the time of rapid storage protein mobilization (seedling length between 20 and 50 mm) showed high proteinase activities in all three assays. Experiments with different inhibitors suggested that at least four enzymes were involved. One of the enzymes resembled mammalian and microbial pepsin-like acid proteinases: the pH optimum was 3.7 and the enzyme was inhibited by pepstatin.—The proteinase activities in the endosperms were high enough to account for the mobilization of the reserve proteins during germination. Moreover the activities at pH 3.7.5.4. and 7.0 in the endosperms were 10-, 25-, and 50-fold the corresponding activities in the growing seedlings (a “reference” tissue). Consequently, it seems that both the acid and neutral proteinases take part in the mobilization of storage proteins in the germinating seed.     

Localization and Activity of a Carboxypeptidase in Germinating Seeds of Scots Pine, Pinus sylvestris

Extracts prepared from the endosperm of germinating seeds of Scots pine, Pinus sylvestris L., hydrolysed two typical carboxypeptidase substrates, Z-Phe-Ala and Z-Phe-Phe, with pH optima at 4.2 and 5.0. The activities were completely destroyed by diisopropylfluorophosphate. Identical heat inactivation curves and elution patterns in gel chromatography on Sephadex G-200 suggest that the two activities are due to a single enzyme. In resting seeds very low carboxypeptidase activity was present in both the endosperm and the embryo. During germination on agar gel at 20°C in the dark the activities, expressed as enzyme units per seed, increased in the seedling and particularly in the endosperm up to the stage when the reserves of the endosperm were completely depleted. The time of rapid increase of activity in the endosperm did not coincide with the onset of storage protein mobilization. On the contrary, the major part of the increase occurred after the bulk of endosperm nitrogen had already been transferred to the seedling. The results suggest that the carboxypeptidase does not play a major role in the mobilization of storage proteins in germinating pine seeds. On the other hand, it probably functions in the proteolytic reactions associated with the senescence of the reserve-depleted endosperm.     

Activities of Two Peptidases in Resting and Germinating Seeds of Scots Pine, Pinus sylvestris

Extracts prepared from resting seeds of Scots pine, Pinus sylvestris L., rapidly hydrolysed two peptides, Leu–Tyr and Ala–Gly, at pH 8.6 and 7.8, respectively. In gel chromatography on Sephadex G-100 the two activities were eluted as separate peaks, which indicates that they are due to two different peptidases. The seeds were allowed to germinate at 20°C, the activities of the two enzymes were assayed separately on extracts from the endosperm and seedling tissues at different stages of germination, and compared with corresponding changes in dry weight and total nitrogen. Both enzyme activities were relatively high in the endosperm of resting seeds, and they increased about 2- and 3-fold during germination (expressed as enzyme units per seed), the increases coinciding with the time of rapid reserve protein mobilization. Both enzymes were also abundant in the embryos of resting seeds, and during germination their activities increased even more rapidly than those in the endosperm. The possible role of these two “alkaline peptidases” in reserve protein hydrolysis is discussed.     

Fire Season Effects on Flowering Characteristics and Germination of Longleaf Pine (Pinus palustris) Savanna Grasses

Pinus palustris (longleaf pine) savannas depend on contiguous grass cover to facilitate frequent surface fires. Enhanced flowering, seed production, and germination may be linked to season of fires. We assessed the effect of month of prescribed fire (February, April, May, and July) on percentage of plants with flowering culms (FCs%), FC/area, FC/plant, seed production, and germination for five warm seasons, fall‐flowering grasses. Multivariate analysis indicated the response of flowering and fruiting to burn month varied among the grass species. The dominant species, Sporobolus junceus and Schizachyrium scoparium var. stoloniferum, had greater numbers of most flowering characteristics when burnt during April, May, and July. Aristida purpurascens had increased FC/plant after May and July burns. In contrast, Ar. mohrii had the fewest FC/plant and seeds/FC when burnt in July. Germination was greatest (26–60%) for Ar. purpurascens. Seeds collected following July burns for Ar. purpurascens and Ar. ternarius and after May burns for S. junceus were within the highest germination values recorded. Germination of Sc. scoparium var. stoloniferum was very low after February and July burns (≤5%). With Ar. mohrii, only seed collected following February (2%) and April burns (3%) germinated. April, May, and July fires increased seed production of dominant matrix grasses, thus facilitating the potential for recruitment of these species and facilitating seed collection from potential donor sites for ground‐layer restoration projects. Varying prescribed fire burn month captured variation in flowering characteristics among these grasses.     

Fire in the Suburbs: Ecological Impacts of Prescribed Fire in Small Remnants of Longleaf Pine (Pinus palustris) Sandhill

Abstract Logging, fire suppression, and urbanization have all contributed to the serious decline and fragmentation of Pinus palustris (longleaf pine) ecosystems in the southeastern United States. Effective management of the remaining patches of these pyrogenic communities must incorporate periodic low‐intensity fires, even where they are located on private lands in populated urban and suburban areas. To explore the effects of fire and its potential use for restoration and management of small fragments surrounded by suburban development, we conducted growing season prescribed fires in remnant longleaf pine sandhill patches in the suburbs of Gainesville, Florida. Density and composition of hardwoods were surveyed pre‐burn and 1 and 9 months post‐burn. Woody stem density decreased in the burn plots, predominantly in the smaller size classes. Flowering responses of forbs and small shrubs were surveyed six times post‐burn for 1 year. Overall, the burns did not yield greater densities of flowering stems, but burn patches had higher species richness and diversity than control patches. In addition, there were consistently greater numbers of “showy flowered” sandhill species in flower in burn patches relative to controls. The results of this research demonstrate that prescribed fire can be used for restoration and management of small remnants of longleaf pine sandhill in suburban neighborhoods. It is also clear that although a single prescribed burn can be effective, it will take more than one burn to attain desired restoration goals in degraded longleaf remnants.     

Light, Seed Coat and Gibberellic Acid in Relation to the Amylase Activity in Germinating Scots Pine Seeds (Pinus silvestris)

The amylolytic activity in homogenates from both embryos and endosperms of Scots pine seeds depends on the presence of α-amylase. The activity of this enzyme has been studied in relation to the light factor during the first 24 hours of the germination process. No positive correlations have been found between the observed activities and earlier observed enhancement of the starch degradation in light. The endosperm activities have been shown to be affected both by the presence of an intact coat and the presence of the embryo. The embryo influence could not be restored by gibberellic acid. The results have been discussed in relation to the physiology of gibberellin and ethylene. The author wishes to express his thanks to the Swedish Natural Science Research Council for financial support and to Miss Brigitte Thierbach for skilful, technical assistance.     

Influence of Light on Germination of Pinus palustris Seeds

Red light with a wavelength of 660 nm promotes germination of longleaf pine (Pinus palustris Mill.) seeds, and far-red light (730 nm) inhibits germination. The promotion-inhibition process is repeatedly reversible, indicating that germination is controlled by the photoreversible reaction of phytochrome. Response varied greatly between single-tree lots and was dependent on the length of time seeds were imbibed at 5°C. Dry seeds did not respond to light treatments when they were subsequently imbibed and tested in darkness. Stratification for 28 days essentially removed the light requirements for germination.     

Response of Six‐Lined Racerunner (Aspidoscelis sexlineata) to Habitat Restoration in Fire‐Suppressed Longleaf Pine (Pinus palustris) Sandhills

Six‐lined racerunner (Aspidoscelis sexlineata) is an indicator species of frequently burned Longleaf pine (Pinus palustris) forests. To evaluate how the species responded to forest restoration, we conducted a mark‐recapture study in formerly fire‐suppressed Longleaf pine forests exposed to prescribed fire or fire surrogates (i.e. mechanical or herbicide‐facilitated hardwood removal) as well as in fire‐suppressed control sites and reference sites, which represented the historic condition. After initial treatment, all sites were exposed to over a decade of prescribed burning with an average return interval of approximately 2 years. We used population‐level response of A. sexlineata as an indicator of the effectiveness of the different treatments in restoring habitat. Specifically, we compared mean numbers of marked adults and juveniles at treatment sites to that of reference sites. After 4 years, restoration objectives were met at sites treated with burning alone and at sites treated with mechanical removal of hardwoods followed by fire. After over 10 years of prescribed burning, restoration objectives were met at all treatments. We conclude that prescribed burning alone was sufficient to restore fire‐suppressed Longleaf pine sandhills for A. sexlineata populations.     

RNA Polymerase II Activity in Developing and Germinating Cowpea Seeds

RNA polymerase II was partially purified from germinating cowpeaseeds and characterized. RNA polymerase II activity per seedwas low soon after flowering but increased thereafter, reachinga maximum after about two weeks. Upon desiccation, the activitydecreased and was very low in quiescent seeds. After germination,the activity increased rapidly. (Received July 23, 1990; Accepted November 30, 1990)     

Asparagine Enhances Starch Accumulation in Developing and Germinating Lupin Seeds

Regulation of starch accumulation in yellow (Lupinus luteus L.), white (L. albus L.), and Andean lupin (L. mutabilis Sweet) developing and germinating seeds was investigated. Research was conducted on cotyledons isolated from developing seeds as well as on organs of germinating seeds, that is, isolated embryo axes, excised cotyledons, and seedling axes and cotyledons. All organs were cultured in vitro for 96 h in different carbon (60 mM sucrose) and nitrogen (35 mM asparagine or 35 mM nitrate) conditions. Ultrastructure observation showed one common pattern of changes in the number and size of starch granules caused by sucrose, asparagine, and nitrate in both developing and germinating seeds. Sucrose increased the number and size of starch granules. Asparagine additionally increased starch accumulation (irrespective of sucrose nutrition) but nitrate had no effect on starch accumulation. Asparagine treatment resulted in a significant decrease in soluble sugar level in all organs of germinating lupin seeds of the three species investigated. The above-mentioned changes were most clearly visible in white lupin organs. In white lupin, starch granules were visible even in cells of sucrose-starved isolated embryo axes where advanced autophagy occurs. The importance of asparagine-increased starch content in the creation of a strong source–sink gradient in developing and germinating lupin seeds is discussed.     

Longleaf pine (Pinus palustris Mill.) fire scars reveal new details of a frequent fire regime

Question: How frequent and variable were fire disturbances in longleaf pine ecosystems? Has the frequency and seasonality of fire events changed during the past few centuries? Location: Kisatchie National Forest, Western Gulf Coastal Plain, longleaf pine–bluestem ecosystem, in relatively rough topography adjacent to the Red River, Louisiana, USA. Methods: Cross‐sections of 19 remnant pines exhibiting 190 fire scars were collected from a 1.2‐km² area. Tree‐rings and fire scars were precisely dated and analysed for the purpose of characterizing past changes in fire and tree growth. Temporal variability in fire occurrence and seasonality was described for the pre‐ and post‐European settlement periods. Seasonality of historic fires was determined by the scar position within the rings. The relationship between fire and drought was investigated using correlation and superposed epoch analysis. Results: The mean fire return interval for the period 1650‐1905 was 2.2 years (range 0.5 to 12 yr). Significant new findings include: evidence for years of biannual burning, temporal variability in fire seasonality, an increase in fire frequency and percentage of trees scarred circa 1790, and synchronous growth suppression and subsequent release of trees coinciding with land‐use changes near the turn of the 20th century. Drought conditions appeared unrelated to the occurrence of fire events or fire seasonality. Conclusions: Multi‐century fire history records from longleaf pine ecosystems are difficult to obtain due to historic land‐use practices and the species high resistance to scarring; however, our results indicate potential for reconstructing detailed fire histories in this ecosystem. Fire scars quantitatively documented one of the most frequent fire regimes known. Fire regime information, such as the temporal variability in fire intervals, prevalence of late‐growing season fire events and biannual burning, provide a new perspective on the dynamics of longleaf pine fire regimes.     

Seed Bank Viability in Disturbed Longleaf Pine Sites

Some of the most species‐rich areas and highest concentrations of threatened and endangered species in the southeastern United States are found in wet savanna and flatwood longleaf pine (Pinus palustris Mill.) communities. Where intensive forestry practices have eliminated much of the natural understory of the longleaf ecosystem, the potential for reestablishment through a seed bank may present a valuable restoration opportunity. Longleaf pine sites converted to loblolly pine plantations and non‐disturbed longleaf sites on the Coastal Plain of North Carolina were examined for seed bank presence and diversity. Conducting vegetation surveys and examining the seed bank using the seedling emergence technique allowed for verification of the seed bank presence, as well as evaluation of the quality of the seed bank on disturbed longleaf pine sites. Forty‐three species and over 1,000 individuals germinated, and the seed banks of both the disturbed and non‐disturbed stand types contained species not noted in the vegetation survey. Although many of these species were considered weedy and typical of disturbance, numerous taxa were indicative of stable longleaf pine communities. This study confirms both the presence and quality of seed banks in highly disturbed former longleaf pine sites, suggesting that the seed bank may be an important tool in restoration efforts.     

Structure and Composition of Historical Longleaf Pine Ecosystems in Mississippi,USA

Longleaf pine (Pinus palustris) historically was a widespread ecosystem composed of a simple tree canopy and grasslands ground layer. After widespread loss of this ecosystem due to logging and fire exclusion, little quantitative information exists about historical structure for restoration goals. We identified composition in De Soto National Forest and Pearl River County, Mississippi, USA, and density, basal area, and percent stocking in Pearl River County using General Land Office surveys and US Forest Service Forest Inventory and Analysis surveys. Historical longleaf ecosystems were about 85% pine, with lesser amounts of broadleaf evergreen and oak species. Densities were about 175 to 180 trees/ha, mean tree diameters were 45 cm, and stocking was around 60% to 65%, which suggested longleaf pines were closed woodlands. Current forests are 38% to 57% pine, primarily loblolly, while longleaf pine is 2% to 8% of composition. Indeed, current longleaf pine composition across the Coastal Plain averages 3% and does not reach 10% at smaller landscape scales. Fire-sensitive broadleaf species of water oak, sweetgum, yellow-poplar, and red maple increased from about 0.5% composition to 2% to 10% of composition. Forests became twice as dense, at about 280 trees/ha to 330 trees/ha, with mean tree diameters of 22 cm. These results characterize conversion from open old growth longleaf forests, resulting in part from human maintenance, to successional forests due to human disruption of the historical ecosystem. It is important to remember structure and composition of historical forests for restoration and recognize wholesale changes so that successional forests do not become the new social and cultural baseline.     

Starch Deposition and Carbohydrase Activities in Developing and Germinating Soya Bean Seeds

Immature soya bean seeds accumulate starch as a transient reservematerial which is utilized later in development. Germinatingseeds also accumulate starch reserves, probably as a resultof gluconeogenesis from storage lipid. Developing beans showa rapid increase in ß-amylase activity which continuesinto early germination before declining. Distribution of ß-amylaseactivity is not consistent with its supposed role in starchdegradation. Soya bean seeds also contain -amylase and -glucosidaseactivities which could be responsible for starch mobilization. Glycine max (L.) Merr., soya bean, starch, carbohydrase, amylase, -glucosidase     

Longleaf Pine (Pinus palustris P. Mill.) Restoration Using Herbicides: Overstory and Understory Vegetation Responses on a Coastal Plain Flatwoods Site in Florida,U.S.A.

A study was conducted on a Coastal Plain flatwoods site in Florida to determine the effects of common forestry herbicides on Longleaf pine seedling survival and growth and on the understory vegetation. Following removal of the overstory slash pine, five low‐rate herbicide treatments were applied over the top of planted Longleaf pine seedlings to provide short‐term understory vegetation control and accelerate seedling growth. The objective was to increase Longleaf pine growth by reducing the shrub competition while increasing the herbaceous ground cover. Despite causing reduction in seedling survival over the control treatment, imazapyr (0.21 ae kg/ha) resulted in the highest seedling growth (height and volume). The significant reduction of shrub cover, density, and height by imazapyr was believed to be responsible for the improved seedling growth in this treatment. Both hexazinone (0.56 ai kg/ha) and sulfometuron methyl (0.26 ai kg/ha) + hexazinone (0.56 ai kg/ha) treatments also reduced cover of Runner oak, a major shrub species, but the response was evident only 8 months after treatment. Although sulfometuron methyl (0.26 ai kg/ha) and sulfometuron methyl + hexazinone treatments did not result in any significant change in overall grass, forb, and shrub cover, both treatments resulted in greater Longleaf pine growth compared to the control. None of the herbicides significantly affected the major understory grasses and forbs. Overall, imazapyr provided the best desired results with significant increase in seedling growth and better control of shrub species with no significant effects on grass and other herbaceous species cover.     

Effects of long-term fire exclusion on tree species composition and stand structure in an old-growth Pinus palustris (Longleaf pine) forest

Frequent fire is an integral component of longleaf pine ecosystems, creating environmental conditions favoring survival and growth of juvenile pines. This study examined stand structure, species composition, and longleaf pine regeneration in an old-growth tract of longleaf pine forest (Boyd Tract) experiencing long-term (>80 yr) fire exclusion in the Sandhills of North Carolina. Sampling of woody stems (i.e., 2.5 cm diameter at breast height) and tallies of longleaf pine seedlings were carried out in plots established randomly on upland, mesic areas and lowland, xeric areas within the Boyd Tract. Dominant woody species in mesic plots were black oak, hickories, and large, sparse longleaf pines. Xeric plots had high densities of turkey oak with the large longleaf pines, as well as higher frequencies of smaller longleaf stems. These differences between areas were associated with higher clay content of upland soils and higher sand content of lowland soils. Age-class frequency distributions for fire-suppressed longleaf pine following the last wildfire at the Boyd Tract approximately 80 yr ago contrasted sharply with data from an old-growth longleaf tract in southern Georgia (Wade Tract) that has been under a long-term frequent fire regime. Post-burn recruitment for the Boyd Tract wildfire appears to have been initially high on both site types. Longleaf pine recruitment diminished sharply on the mesic site, but remained high for 60 yr on the xeric site. Currently, longleaf pine regeneration is minimal on both site types; several plots contained no seedlings. Sharp contrasts in longleaf pine dominance and stand structure between the Boyd and Wade Tracts demonstrate the importance of large-scale disturbance, especially hurricanes and fire, in shaping the structure and function of longleaf pine ecosystems of the southeastern United States. In particular, long-term exclusion of fire on the Boyd Tract has altered stand structure dramatically by permitting hardwoods to occupy at high densities the characteristically large gaps between longleaf stems that are maintained by fire and other disturbances.