Influence of high temperatures on seed germination of a special Pinus pinaster stand adapted to frequent fires

This study examines the effect of fire on the germination of Pinus pinaster seeds from a population with special adaptations to this type of disturbance, due to the high frequency of fires to which they have been subjected. The action of fire was simulated in the laboratory using thermal shocks. Temperatures of 60, 90, 120, 200 and 300 °C were used for exposure times of 1 and 5 min. The viability of seeds of this species from the soil seed bank of a population subjected to a wildfire was also evaluated. The results show that germination is not increase by a thermal treatment. The effect is negative at temperatures above 90 °C and exposure times of 1 min with a significant decrease in, or even no, germination. Therefore the seeds cannot stand high temperatures and reproductive effort is placed on serotinous cone development and the production of high numbers of seeds. The high viability of the seeds from the area burned by wildfire, immediately after the fire and 1 year later, show the high potential of the soil seed bank to contribute to the regeneration of the burned area.     

Genetic diversity and differentiation of two Mediterranean pines (Pinus halepensis Mill. and Pinus pinaster Ait.) along a latitudinal cline using chloroplast microsatellite markers

Several studies have reported glacial refugia and migration pathways for different pine species in the Iberian Peninsula, all of them based on a single‐species approach. In this paper, chloroplast microsatellites (cpSSRs) are used to compare population genetic structure and diversity estimates for interspecific pairs of populations located along a cline from southwestern (latitude 36°32′ N, longitude 5°17′ W) to northeastern Spain (latitude 42°14′ N, longitude 2°47′ E) in two widely distributed Mediterranean pines, Pinus halepensis Mill. and Pinus pinaster Ait. Some cpSSRs were shared between species, facilitating comparison of levels of gene diversity at the species level and inferences about within and among species differentiation. P. pinaster showed a much higher number of variants (29) and haplotypes (69) than P. halepensis (20 and 21, respectively). Moreover, genetic diversity estimates for interspecific pairs of populations along the cline were negatively correlated. Three main causes may explain the differences between species in the present‐day distribution of genetic diversity: (1) the distribution of genetic variability before the Quaternary glaciations, with an earlier presence of P. pinaster in the Iberian Peninsula and a late spread of P. halepensis from eastern and central Europe, (2) the location of the Holocene glacial refugia and the migration pathways from these refugia to the present‐day range (from northeast to southwest in P. halepensis and from southwest to northeast and northwest in P. pinaster) and (3) the interactions between species during the postglacial spread.     

Seasonal fluctuations in soil phenolics of a coniferous forest: effects on seed germination of different coniferous species

This study examined seasonal variations in the content of water-soluble phenolics extracted from litter and Ah horizon of forest soil under Pinus laricio Poiret trees. After having identified and quantified different phenolics, the effects of seasonal phenolic extracts, single phenolic acids and synthetic phenol mixtures on seed germination of Pinus laricio Poiret, Pinus pinaster Aiton and Pinus halepensis Mill. were evaluated, and the activity of phenolics on the main enzymes involved in the glyoxylate cycle was tested. The results indicated a seasonal variation of phenol content in soil, with largest concentrations of water-soluble phenolic acids in autumn and smallest concentrations in summer. Addition of phenol extracts to germination medium reduced seed germination showing phytotoxic effects which differed, depending on the species and the fractions tested. Phenols extracted from litter and Ah horizon in autumn and winter, vanillic acid, and synthetic mixtures reproducing phenols extracted from soil in winter inhibited seed germination of Pinus laricio, pinaster and halepensis more than other treatments. A greater inhibitory effect was observed on seed germination of Pinus pinaster and halepensis compared to Pinus laricio. The phenols also had the greatest inhibitory effect on glyoxylic enzyme activities. Section Editor: H. Lambers     

Effect of high temperatures on seed germination of two woody Leguminosae

Cytisus scoparius and Genista florida regenerate after fire by stump-sprouting but also by seed. Seeds of these species were heated to a range of temperatures similar to those registered on the surface soil during natural fires (from 50 to 150 °C) and a range of exposure times (from 1 to 15 min). No germination was observed at high temperatures, 130 °C, when the exposure time was 5 min or more. However, moderate heat treatments (at 70 and 100 °C) significantly increased the rate of germination relative to controls. Cytisus scoparius is more favoured by fire action than Genista florida, with germination rates slightly greater following 100 °C for 5 min and 130 °C for 1 min than after mechanical scarification.     

Influence of heat on seed germination of seven Mediterranean Leguminosae species

The influence of high temperatures (dry heat and hot water) on germination of seven Mediterranean Leguminosae species typical of fire-prone ecosystems in southern Spain is analyzed, in order to know the response of seeds to wildfires and the possible implications in their regeneration after this disturbance. Seeds were heated to a range of temperatures (50 °–150 °C) and exposure times (1–60 min) similar to those registered in the upper soil layers during wildfires. Germination tests were carried out in plastic Petri dishes over 60 days. In general, the degree of seed germination promotion by dry heat treatments showed a wide interspecific variation, although the final germination level was increased in all the studied species except for Scorpiurus muricatus. The thermal pretreatment of 50 °C, however, was not effective for germination in any species, and rising the temperature to 70 °C only slightly enhanced the germination in Cytisus patens. The preheatings of 90 °C (5 and 10 min), 120 °C (5 and 10 min), and 150 °C (1 min) were the most effective in promoting seed germination. Hot water (100 °C) scarification also increased the final germination level in all cases, with the exception of C. patens. The germination rates after preheating were much lower than in mechanically scarified seeds and closely resembled those of the untreated seeds, except for C. reverchonii, whose seed germination rate decreased with heat. The response of species to heat shock had no clear relationship with life trait or with the specific post-fire regeneration strategy (obligate seeder or facultative resprouter). Those species coexisting in the same habitats had different heat optimal requirements for seed germination, an strategy suggested by some authors as minimizing interspecific competition in the secondary succession started after fire.     

Heat effects on seeds and rhizomes of a selection of boreal forest plants and potential reaction to fire

To analyse the potential reaction to firegenerated heat pulses, seeds of 12 species of plants and rhizomes of three species were exposed to elevated temperatures for 10 min. The tested material split into three groups with respect to heat tolerance: (1) the rhizomes, for which the lethal temperatures were in the range 55–59° C; (2) the seeds of most of the species tested, for which the lethal temperatures were in the range 65–75° C; (3) The seeds of two species of Leguminosae and three species of Geranium for which the lethal temperatures were around 100° C. For all three Geranium species and for one of the legume species, Anthyllis vulneraria, exposure temperatures above ca. 45° C resulted in dormancy release, and maximum germination occurred above 60–65° C. Speed of germination was little affected for most species, except after exposure to nearlethal temperatures, where it slowed down dramatically, although the seedlings emerging were healthy. We conclude that due to sharp temperature gradients in the soil during fire, differences in heat tolerance between species in most cases are not large enough to be a decisive factor in their post-fire colonising success. There are exceptions: the seeds of certain taxa that are impermeable to water in the dormant state, some of which have heat triggered germination.     

The effect of high temperatures on seed germination of one native and two introduced conifers in Patagonia

We examined the effect of thermal shock on the germination of seeds of three conifers, two introduced (Pseudotsuga menziesii and Pinus ponderosa), and one native to Patagonia (Araucaria araucana). Previous research has suggested increased susceptibility to invasions in burnt areas, and therefore, the effect of simulated fire (heat) on seed germination in these native and introduced species was compared. Seeds were heated to two different heat intensities (50°C and 100°C) for 1 or 5 min, which is within the temperature range reached in the upper soil layers during forest fires. Germination tests were then carried out in a growth chamber. The heat treatments had a negative effect on the germination of P. menziesii at temperatures of 100°C, and a negative effect on the germination of P. ponderosa at the temperature of 100°C and the exposure of 5 min. The heat treatments had no affect at all on A. araucana. The species with larger seeds (A. araucana) had higher survival rates after the thermal shocks. Also intraspecific differences in seed sizes possibly point at larger seeds surviving thermal shocks better than smaller seeds. In addition, thermal shock caused a delay in the onset of germination in the two introduced species, while it did not change the time for germination in A. araucana.     

Pinus halepensis,Pinus pinaster,Pinus pinea and Pinus sylvestris Essential Oils Chemotypes and Monoterpene Hydrocarbon Enantiomers,before and after Inoculation with the Pinewood Nematode Bursaphelenchus xylophilus

Pinewood nematode (PWN), Bursaphelenchus xylophilus, is the causal agent of pine wilt disease, a serious threat to global forest populations of conifers, especially Pinus spp. A time‐course study of the essential oils (EOs) of 2‐year‐old Pinus halepensis, Pinus pinaster, Pinus pinea and Pinus sylvestris following inoculation with the PWN was performed. The constitutive and nematode inoculation induced EOs components were analyzed at both the wounding or inoculation areas and at the whole plant level. The enantiomeric ratio of optically active main EOs components was also evaluated. External symptoms of infection were observed only in P. pinaster and P. sylvestris 21 and 15 days after inoculation, respectively. The EO composition analysis of uninoculated and unwounded plants revealed the occurrence of chemotypes for P. pinaster, P. halepensis and P. sylvestris, whereas P. pinea showed a homogenous EO composition. When whole plants were evaluated for EO and monoterpene hydrocarbon enantiomeric chemical composition, no relevant qualitative and quantitative differences were found. Instead, EO analysis of inoculated and uninoculated wounded areas revealed an increase of sesquiterpenes and diterpenic compounds, especially in P. pinea and P. halepensis, comparatively to healthy whole plants EOs.     

Ectomycorrhizal fungi of Pinus pinaster

A study was undertaken to determine the ability to form ectomycorrhizae with Pinus pinaster Ait. in pure culture syntheses of 98 isolates of putative mycorrhizal fungi, mainly collected in northern Spain. A total of 35 species in 16 genera — Amanita, Cenococcum, Collybia, Cortinarius, Hebeloma, Laccaria, Lactarius, Lyophyllum, Melanogaster, Paxillus, Pisolithus, Rhizopogon, Scleroderma, Suillus, Thelephora and Xerocomus — formed ectomycorrhizae. Many of these fungal species were not previously reported as symbiotic with Pinus pinaster. Results obtained increase the range of potential fungal candidates for inoculation of nursery seedlings.     

Effect of high temperature on moisture loss,imbibition and germination of seeds ofTrianthema Govindia Buch.-Ham. ex DC

Summary Seeds of erect and prostrate plants ofTrianthema govindia Buch. ham. ex DC., growing in shade and open respectively, differed significantly in seed weight and percentage germination. Effect of high temperature exposure to these seeds has been studied in view of water depletion, imbibition and seed germination. The seeds of both the types were subjected to temperatures of 40, 50, 60, and 70° C for 24, 48, 96, and 144 hours. The three factors viz., loss of water, water imbibition and germination of seeds were positively correlated to the duration of treatment at different temperatures. A highly significant positive correlation was also observed between moisture depletion and imbibition, and between imbibition and germination. The percentage germination was favoured at 40° C in both the types of seeds and was increased with the increase of treatment duration. However, at higher temperatures (50 and 60° C) the percentage declined while at 70° C the seeds lost their vitality.     

Influence of temperature upon growth and sporulation in two species of Phytophthora

The paper deals with the influence of temperature on the growth and sporulation of two species ofPhytophthora, viz.,P. palmivora Butl. andP. parasitica Dast. var.macrospora Ashby, the causal agents of fruit rots ofAchras sapota L. andAnona squamosa L. respectively. Germination of sporangia at different temperatures were also undertaken. There was marked variation in growth and sporulation of these two organisms. Isolate C (Phytophthora palmivora) showed no growth at 5° and 35°C, scanty growth at 10° and 32.5° with an optimum temperature between 26–28°C. On the other hand, Isolate S (Phytophthora parasitica var.macroscora) showed no growth at 10°C, but slight growth even at 37°C. Eight days exposure at 37°C completely stopped the growth of this Isolate. It showed best growth at 30°C and hence this was its optimum temperature. In general, Isolate C sporulated abundantly at all temperatures tested but reached its maximum at 25°C. On the other hand Isolate S showed best growth but failed to sporulate at any of the temperatures in 98 hours growth, although it sporulated freely when the incubation period extended up to two weeks. On the basis of temperature toleration the twoPhytophthora isolates are distinguished from each other as two different species. This confirms the earlier observations and nomenclature criterion as emphasized and formulated byTucker (1931). In the germination studies, it was observed that the indirect germination with the formation of abundant zoospores started from 5° and continued even up to 35°C, reaching maximum at 20°C. High temperature was not favourable for indirect germination. As the temperature proceeded increasing, the percentage of direct germination by formation of germ tubes also increased. Direct germination was observed from 10° which continued up to 37°C, with a maximum reach at 30°C. This confirms the epidemic of fruit rots in nature during monsoon season which is prevalent with the persistence of high humidity and rainfall.Taken from a thesis submitted by the author for the degree of Master of Science in the Faculty of Agriculture, Poona University, India.     

Ex vivo determination of potentially virulent Sporothrix schenckii

Hyphae from 30 isolants ofSporothrix andOphiostoma species were washed, dried and pyrolyzed at 350°C. Pyrolysis products were separated on a Carbowax column heated 7.5°C/min to and maintained for 50 min at 160°C. Hydrogen flame detector responses were recorded graphically. Fifteen clinical isolants ofS. schenckii from geographically separated sources produced qualitatively identical pyrograms.S. foliorum, 8 avirulentS. schenckii and otherSporothrix species isolants from soils, andSporothrix states of 6Ophiostoma species yielded pyrograms readily distinguished from each other and from those of virulentS. schenckii. Taxonomic and clinical implications of the pyrograms are mentioned.     

Constraints on tree seedling establishment after fires: passing the germination bottlenecks

• Persistence and colonization by tree species in an environment following a fire depends on the effects on seed germination and seedling development. We used seeds of Kielmeyera coriacea and Qualea parviflora as a model to test the effects of high temperatures on germination and initial development of tree seedlings.
• We exposed the seeds to heat flow (70, 100, 130, 150 or 170 °C) for 2 or 5 min and compared the germination with that of unheated seeds (control). Seedlings were then harvested after 3, 7 or 15 days to evaluate aerial and root mass, root:shoot ratio, presence of cotyledon opening, true leaves, and secondary roots.
• We found no effect on germination for seeds exposed to temperatures ≥150 °C. However, germination was significantly reduced for seeds exposed to 100 °C for both 2 and 5 min. The mass of 15-day-old K. coriacea seedlings was smaller when seeds were heated at 70 °C for 5 min or at temperatures higher or equal to 100 °C. Qualea parviflora seedlings did not show any difference in mass, but there were marginal differences in the presence of roots and the opening of cotyledons. Kielmeyera coriacea seedlings allocated biomass faster than Q. parviflora.
• High temperatures affect both quantity and quality of germinable seeds, as well as biomass allocation during initial seedling development. These factors may explain the decrease in seedlings observed after fire, suggesting a bottleneck effect that influences population dynamics and species persistence in systems with frequent fires.

     

Germination response to fire-related factors of seeds from non-serotinous and serotinous cones

Pinus halepensis, a Mediterranean pine tree, is a partially serotinous species: individual trees of this species carry both non-serotinous and serotinous cones. Serotinous cones open mainly after fire, whereas non-serotinous cones open in absence of fire. In this study we addressed the question, whether or not this cone response is linked with the germination response of seeds to fires. Two main factors associated with fire are heating of seeds and soil pH. A combination of high heat and high pH simulates a scenario with fire, whereas low heat and low pH simulates a scenario without fire. We assessed the separate and combined effects of heat and pH on the germination rate and the percentage of germination of seeds from non-serotinous cones and two age classes of serotinous cones of P. halepensis. Heat had no effect on the percentage of germination of seeds from any of the cone types, but did positively affect the germination rates of seeds from both age-classes of serotinous cones. High pH negatively affected the germination rate of seeds from all cone types as well as the percentage of germination of seeds from non-serotinous cones. The combinations of heat and pH had different effects on the three cone types: percentage of germination and rate of germination of seeds from non-serotinous cones was higher in the combination high heat-high pH than in the combination low heat-low pH. In the combination high heat-high pH, seeds from serotinous cones germinated better than seeds from non-serotinous cones. The different germination responses of seeds from non-serotinous and serotinous cones could not be attributed to differences in cone age. Our results indicate that the cone response is linked to the germination response of the seeds in P. halepensis, with seeds from serotinous cones being more tolerant to fire related factors. This revised version was published online in June 2006 with corrections to the Cover Date.     

Temperature effects on dormancy levels and germination in temperate forest sedges (Carex)

The effects of stratification temperatures and burial in soil on dormancy levels of Carex pendula L. and C. remota L., two spring-germinating perennials occurring in moist forests, were investigated. Seeds buried for 34 months outdoors, and seeds stratified in the laboratory at temperatures between 3 and 18 °C for periods between 2 and 28 weeks, were tested over a range of temperatures. Seeds of the two species responded similarly to stratification treatments, except for an absolute light requirement in C. pendula. Primary dormancy was alleviated at all stratification temperatures, but low temperatures were more effective than higher ones . (≥ 12 °C). Dormancy induction in non-dormant seeds kept at 5 °C occurred when seeds were subsequently exposed to 18 °C. Dormancy was not induced by a transfer to lower temperatures. Buried seeds of both species exhibited seasonal dormancy cycles with high germination from autumn to spring and low germination during summer. Temperatures at which the processes of dormancy relief and of dormancy induction occurred, overlapped to a high degree. Whether, and when, dormancy changes occurred depended on test conditions. The lower temperature limit for germination (> 10%) was 9 °C in C. remota and 15 °C in C. pendula. Germination ceased abruptly above 36 °C. Germination requirements and dormancy patterns suggest regeneration from seed in late spring and summer at disturbed, open sites (forest gaps) and the capability to form long, persistent seed banks in both species.     

Lokalrassen und dreinadelige Kurztriebe bei der Krummholz-Kiefer (Pinus mugo) auf der Rax-Alpe (Niederösterreich)

On the Rax-Alpe in Lower AustriaPinus mugo occurs in four local races which differ in colour and length of male flowers, and cone symmetry. On the thicker branch bases of one plant three-needled besides the ordinary two-needled fascicles were found. This phenomenon also occurs in the MediterraneanP. pinea, P. pinaster andP. halepensis, and may represent a common primitive trait.     

Dormancy inDioscorea: Differences of temperature responses in seed germination among six Japanese species

Effects on seed germination of temperatures ranging from −2 C to +29 C were tested inDioscorea nipponica, D. tokoro, D. japonica, D. tenuipes, D. septemloba andD. quinqueloba which orginate in the temperate zone; they are distributed from northern cold areas to southern warm areas approximately in this order in Japan. After water imbibition of these seeds, chilling induced full germination, and high temperatures over 23 C induced a secondary dormancy, but sensitivities to the chilling and to the high temperatures differed with species. Cold-climate species germinated rapidly at higher temperatures after a short-term chilling or even without chilling, whereas warm-climate species required chilling of a rather long period for germination; thus, among 6 species tested, favourable temperatures for germination and climatic temperatures of distribution area were conversely correlated. Seeds ofD. tokoro andD. japonica collected from several populations grown in different climates were also tested for germination at 11 to 29 C; seeds from warm climates germinated rather slowly compared to seeds from cold climates. These inte- and intra-specific adaptation manners in the temperature members of the genusDioscorea are entirely different from those of many other plant genera reported by some workers.     

Effects of forest fire ash on germination and early growth of four pinus species

Fire modifies the germination of seeds of numerous species. One of the fire factors that cause these modifications is the ash. This study analysed the germination of seeds of Pinus sylvestris L., Pinus nigra Arn. Pinus radiata D. Don and Pinus pinaster Aiton, subjected to different ash treatments, and the development of seedlings that grew in these conditions for 14 weeks. We obtained the ash by completely burning leaves and small twigs from the most abundant woody species in populations of pines. The ash treatments applied were as follows: Control (without ash), Low (half of the amount registered in a fire), Medium (equal to the amount registered in a fire) and High (double the amount registered in a fire). Germination took place on paper inside Petri dishes and in soil, and growth was only analysed for the plants that grew in the soil. For all species, and especially P. nigra, germination rate decreased as the amount of ash applied increased. The negative effects of the ash were more apparent following treatments in Petri dishes than in the soil. In the dishes, the average germination time varies little between the four species of pine. In soil, the average germination time is more prolonged, and only some differences were observed between treatments in P. sylvestris and P. nigra. In these species the high treatment significantly increased the average germination time. We found a greater sensitivity of P. sylvestris and P. nigra to the addition of ash, showing a higher seedling mortality rate. However, it appears that seedling development (length and weight) was not affected in any of the species through the addition of ash. These tests allow us to deduce that, in P. sylvestris, P. nigra, P. radiata and P. pinaster, the ash produced by forest fires has an inhibiting effect on germination and little effect on the development of seedlings in the first months of life. This revised version was published online in June 2006 with corrections to the Cover Date.     

Distribution and sporulation phenology of myxomycetes in the Sonoran Desert of Arizona

All pith samples from 68 dead saguaro cacti in 3 plots and 11 isolated dead plants in Saguaro National Monument, Arizona, produced at least one species of myxomycete upon incubation at 20 or 30°C. Three species,Badhamia gracilis (Macbr.) Macbr.,Physarum straminipes Lister, andDidymium eremophilum M. Blackwell et Gilbertson, developed at high frequencies on the substrates in moist chamber culture.Perichaena corticalis (Batsch) Rost, andProtophysarum phloiogenum M. Blackwell et Alexopoulos were also present. Although previous literature reports [9] indicated that Myxomycetes grow best at low pH, these species all tolerated substrates of pH 8.7–10.4.Didymium eremophilum andP. phloiogenum had peaks in sporulation within 6 days; other species were slower. There was no difference in time of sporulation ofB. gracilis orD. eremophilum at 20 and 30°C; however, sporulation ofP. straminipes was significantly later at 30°C. Reduced spore germination and slower buildup of critically sized amoebal populations ofP. straminipes at 30°C may be a factor.     

The maternal environment determines the timing of germination in Pinus pinaster

Optimizing the germination timing is crucial for the establishment of new generations. We hypothesized that environmental maternal effects may be relevant in the fine tuning of this trait in a long-lived Mediterranean model tree. We analyzed the influence of maternal genotype, maternal environment and their interaction on the germination success and germination phenology of 8725 Pinus pinaster seeds collected from genotypes clonally replicated in two contrasting environments. Besides maternal genetic effects, the maternal environment significantly affected both the percentage and the timing of germination. Seeds from the more favourable environment germinated 7.5 days earlier and showed higher germination rate (0.93 ± 0.01 vs 0.85 ± 0.03). Seed weight significantly influenced germination time, but seed weight differences between maternal environments were not enough to explain this form of transgenerational plasticity. The effect of the maternal environment varied depending on the genotype, indicating that genetic variation in the sensitivity to the maternal environment in this pine species does exist.