Blue stain fungi associated withTomicus piniperda (Coleoptera: Scolytidae) on Japanese red pine

A number of various species of blue-stain fungi were isolated fromTomicus piniperda adults at various stages of development, as well as from the galleries, pupal chambers and sapwood underneath galleries on Japanese red pine. This study was an attempt to identify the species, composition of blue-stain fungi associated withT. piniperda, the frequency of occurrence of the fungi, and their role in the sapwood-staining of Japanese red pine in Tsukuba City, central Japan. Among the seven species of blue-stain fungi isolated, an undescribed species ofOphiostoma together withO. minus were the dominant species and closely associated withT. piniperda. These two species occurred on newly emerging adults more frequently than the overwintered adults.Hormonema dematioides was also associated with the beetle, however, its frequency of occurrence from the emerged new adults was very low. Although the two other species,O. ips andGraphium sp. were also isolated from emerged beetles, the frequency of these fungi from gallery systems suggested that they were accidentally carried byT. piniperda. Leptographium wingfieldii, known to be associated with the beetle in Europe, was also isolated at a very low frequency and the fungus seemed not to be closely associated with the beetle.Ophiostoma sp. andO. minus appear to be the most important causes of blue-stain of Japanese red pine sapwood after infestation byT. piniperda.     

Comparison of organic matter dynamics in soil between Japanese cedar (Cryptomeria japonica) forest and adjacent Japanese red pine (Pinus densiflora) forest established on flatland

In order to clarify the effects of tree species on organic matter dynamics in soil, we investigated the amount of forest floor material, leaf litter decomposition rate, soil chemical characteristics, soil respiration rate and cellulose decomposition rate in a Japanese cedar forest (cedar plot) and an adjacent Japanese red pine forest (pine plot) established on a flatland. The amount of forest floor material in the cedar plot was 34.5 Mg ha⁻¹ which was greater than that in the pine plot. Because the leaf litter decomposition rate was higher in the pine plot than in the cedar plot, it is likely that the difference in the amount of forest floor material between the plots is caused by the difference in the leaf litter decomposition rate. The C concentrations of soil in the cedar plot were 1.2–2.1 times higher than those in the pine plot. Soil pH(H₂O)s in the cedar plot were significantly higher than those in the pine plot. The soil respiration rates and the rates of mineralized C in the cedar plot byin vitro incubation were higher than those in the pine plot. From this result, it is assumed that soil organic matter in the cedar plot was decomposed relatively faster compared with the pine plot. Furthermore, microbial activities, which were reflected as cellulose decomposition rates in the cedar plot, were higher than those in the pine plot. A part of this paper was presented at the 109th Annual Meeting of the Japanese Forestry Society (1998).     

Response of Japanese red pine to inoculation with a blue stain fungus,Ceratocystis piceae

To confirm the pathogenicity of a blue stain fungus,Ceratocystis piceae (Münch) Bakshi to the Japanese red pine (Pinus densiflora Sieb. et Zucc.), the responses of healthy young pine trees and stressed trees which were girdled by the half-circumferential girdling technique were investigated by the fungal inoculation test. Although neither of the pine trees inoculated withC. piceae in the non-girdled treatment nor the controls died, mortality of the trees girdled and inoculated withC. piceae was 28.6%. In the pine trees inoculated withC. piceae, the mean area of the necrotic lesion of the sapwood was significantly larger than that of the controls, and the mean of the water pressure potential of the xylem decreased, regardless of the girdling treatment. TheC. piceae was reisolated from the wood pieces near the inoculation points on the inoculated trees, but not from the controls. These results suggest that under strongly stressed conditions, the Japanese red pine trees might have been killed by heavy infestations ofC. piceae carried by bark beetles. A part of this paper was presented at the 103rd Annual Meeting of the Japanese Forestry Society (1992).     

The role of litter and humus layer as a mortality factor of seedlings of Japanese red pine,Pinus densiflora Sieb. et Zucc.

The process of decrease in seedlings of Japanese red pine (Pinus densiflora Sieb.et Zucc.) and the effect of litter and humus layers on the mortality were studied in their stand and adjacent bare land. After the rainy season, only small number of individuals died in the bare land but more seedlings died in the forest. The number of seedlings killed by drought was greater in the forest than in the bare land. The population of naturally occurring current year seedlings was larger in the forest than in the bare land, but that of 1- to 5-year-old seedlings was smaller. Severe summer drought exceeding the permanent wilting point was observed only in the forest F-H horizon where greater part of seedling roots distributed, this may be one of major mortality factors. Inhibitory substances to seedling growth in fresh fallen pine needles were also suggested.     

Photosynthesis and chlorophylla fluorescence of two poplars under water stress

In order to estimate drought tolerance in two species ofPopulus, Populus alba var. pyramidalis Bunge andPopulus nigra L. var. thevestina (Dode), widely planted at the southern margin of the Taklimakan Desert, responses of net photosynthesis and chlorophylla fluorescence to irradiance and water stress were examined under laboratory conditions. Results showed thatP. alba exhibited stronger drought tolerance thanP. nigra. A linear relationship between net photosynthetic rates (A _n) and electron transport rates (ETR) was found in both poplars under different irradiance and leaf water potentials. Net photosynthetic rates (A _n) in the two poplars significantly correlated linearly with the photochemical efficiency of the saturation light-adapted leaves throughout the range of leaf water potentials, suggesting that the leaf photochemical efficiency in saturation light-adapted leaves can be used to estimate leaf photosynthetic capacity and leaf water conditions in the two poplars within a magnitude of air temperature between 20 and 30°C. This research was supported by the Japan Society for the Promotion of Science (CAS-9601), the Chinese Academy of Sciences and partly supported by the National Natural Foundation of China (39870154).     

Lateral female strobili production in a Japanese red pine (Pinus densiflora Sieb. et Zucc.) clone by exogenous cytokinin application

Application ofN ⁶-benzylaminopurine (BAP) and gibberellin (GA₃) to a grafted Japanese red pine (Pinus densiflora Sieb. et Zucc.) clone promoted flowering. BAP (250 mg/l) was sprayed five times to the top of branches in July or September 1992. GA₃ (100 or 500 mg/l) was applied five times to the whole plants by spray treatment during the above period, or GA₃ solution of the same concentration was injected once into a small incision made in the main stem of clone trees at the beginning of July or the middle of September 1992. In May 1993, no apical female strobili were observed at the top of the new shoots in any treatment. However, lateral female and bisexual strobili were produced in the lower part of new shoots by BAP application in September, regardless of GA₃ application. BAP applied in September without GA₃ application was sufficient to produce lateral female and bisexual strobili. In such treatment, production of female and bisexual strobili was 47 and 25, and they occurred in 16 out of 68 new shoots, 11 out of 15 treated branches and 5 out of 5 treated trees. The average number of female and bisexual strobili per new shoot was 1.06. BAP treatment either in July or in September significantly decreased (p<0.05) the production of male strobili. These results indicate that BAP application induces the development of lateral female strobili in Japanese red pine when applied during the period of floral differentiation. A part of this paper was presented at the 105th annual meeting of the Japanese Forestry Society.     

A comparison of needle retention on Japanese black pine and Japanese red pine

Needle retention (number of needle sets), needle density, height increment and radial increment were surveyed on seven Japanese black pines (Pinus thunbergii Parl.) and seven Japanese red pines (Pinus densiflora Sieb. et Zucc.) growing on the same site. Number of needle sets on branches with respect to whorl position was estimated visually. In 1999, maximum summer needle retention was observed on the fifth and sixth whorl from the top, with values of 3.4 needle sets for the black pines and 2.4 needle sets for the red pines, respectively. The needle trace method (NTM) was used to determine needle retention and needle density along the main stems retrospectively for the years 1968–1998. The long-term mean summer needle retention along the main stem was 3.7 needle sets for the black pines and 2.2 needle sets for the red pines. In both pine species, the number of needle sets varied from year to year. However, the long-term budget between newly born and annually shed needle sets was in equilibrium. The long-term average of needle density per cm of stem shoot was 9.4 needle pairs for the black pines and 7.4 needle pairs for the red pines. The results showed clear intra-specific and inter-specific similarities in needle retention, height increment and radial increment trends. The values for number of needle sets, height increment and radial increment positively increased with favorable growing conditions, whereas the value of needle density had an opposite tendency.     

Effect of electric impulse on sporocarp formation of ectomycorrhizal fungusLaccaria laccata in Japanese red pine plantation

We examined the effect of electric impulses on sporocarp formations of ectomycorrhizal fungus in a Japanese red pine (Pinus densiflora) plantation.Laccaria laccata was the dominant species in this study plantation. Spatial distribution concentrated in the electric impulse treatment zone, and recorded a markedly larger number of sporocarps compared with untreated control plots. The effect of electric impulses was also recognized onL. laccata fruiting with a mycorrhizal sapling in a pot. Sporocarp production ofL. laccata was strongly promoted by the electric impulse in experiments bothin vivo natural plantation stands andin vitro pot saplings. Our work indicates the possibility of enhancement of sporocarp production of the mycorrhizal fungi with electric impulses in the nursery and plantation. This work was partly supported by a grant from the Kyushu Electric Power Company.     

Summer field performance of <Emphasis Type="Italic">Quercus petraea</Emphasis> (Matt.) Liebl and <Emphasis Type="Italic">Quercus pyrenaica</Emphasis> Willd seedlings,planted in three sites with contrasting canopy cover

In 2000, one-year-old seedlings of pyrenean oak (Quercus pyrenaica Willd.) and sessile oak (Quercus petraea [Matt.] Liebl) were planted in a thinned and an unthinned plot in a pinewood (Pinus sylvestris), and in a nearby clearing. In summer 2002 and 2003, water relations and gas exchange parameters were measured to address the impact of drought on the seedlings. Chlorophyll a fluorescence was also measured to explore leaf photochemistry and a possible non-stomatal limitation to photosynthesis (A). Reduction in stomatal conductance (g) in response to the decrease of predawn water potential (Ψ_pd) resulted the main cause affecting net carbon uptake. Water potential at midday (Ψ_md) was similar in both species but Quercus petraea was more sensitive to soil water deployment occurred along summer, showing slightly lower Ψ_pd because worse recover of water potential during night. Rate of photosynthesis was higher in Q.␣pyrenaica probably in relation to its greater leaf mass per area (LMA) and nitrogen content per leaf area (N_a). Mortality was highest in the clearing and lowest in the thinned pinewood. Throughout the summer, soil moisture was higher in the thinned area, possibly because of the reduction in tree transpiring surface and interception of rainfall. Accordingly, Ψ_pd of both species was higher in the thinned site.     

Recessive inheritance of morphological characteristics of utsukushimatsu,<Emphasis Type="Italic">Pinus densiflora</Emphasis> f.<Emphasis Type="Italic">umbraculifera</Emphasis>

Pinus densiflora f.umbraculifera, commonly known as utsukushimatsu, is a distinctively shaped form of Japanese red pine whose growth is restricted to a forest stand in Shiga Prefecture, Japan. The inheritance mode of morphological characteristics of utsukushimatsu was studied to preserve the genetic resource of this pine. As previously reported, F₁ trees grown from open-pollinated seeds harvested from trees inhabiting the native stand showed two phenotypes: one resembling utsukushimatsu, which produces multiple trunks, and the other resembling normalP. densiflora, which produces one or a few trunks. In the present study, controlled pollination was carried out using F₁ and normalP. densiflora trees. Segregation ratios of the two phenotypes observed in the F₂ population showed that the morphological characteristics of utsukushimatsu are inherited recessively. This suggests that the mutation of one gene or a few closely linked genes controls the morphological characteristics of utsukushimatsu. Since multiple trunk formation of utsukushimatsu might be related to a loss of lateral bud inhibition, it follows that a simple gene mutation breaks apical dominance inP. densiflora.     

干旱处理对毛竹光响应的影响:基于4种模型比较分析

[目的]采用4种模型对干旱处理下毛竹叶片光响应曲线进行拟合比较,分析各模型的优缺点,以选出适合毛竹光响应曲线拟合的最佳模型,并量化干旱处理对毛竹光合光响应的影响。[方法]主要采用Li-6400便携式光合仪测定干旱处理下毛竹的光响应曲线,并通过4种模型对其进行拟合。[结果]研究表明:(1)在干旱和对照处理下各模型的决定系数R~2均大于0.995,说明拟合精度较高。综合各参数可以看出,直角双曲线修正模型拟合效果最佳,相对误差RE平均值为0.07,而指数模型参数拟合效果最差,RE平均值为0.767;(2)通过直角双曲线修正模型拟合各光合参数,在干旱环境下,暗呼吸速率R_d、光补偿点LCP和水分利用效率WUE较高,分别为对照环境的1.57倍、1.66倍和1.06倍,而净光合速率P_(nmax)、光饱和点LSP和表观量子效率AQY分别比对照环境降低3.6%、17.6%和4.8%。[结论]从整体拟合效果来看,这4种模型拟合顺序为:直角双曲线修正模型直角双曲线模型非直角双曲线模型指数模型。通过在干旱和对照间毛竹各光合参数的比较发现,毛竹对干旱环境有一定的适应能力。     

Growth responses of red pine seedlings to the chemical bioregulator,DCPTA

The chemical bioregulator, 2-(3, 4-dichlorophenoxy)-triethylamine (DCPTA), which has markedly stimulated photosynthesis, carbon allocation, and above- and below-ground growth of several agricultural crops, caused small growth effects on red pine (Pinus resinosa L.) germinants and one and two-year-old seedlings, mainly to foliage and stems. At low DCPTA concentrations, i.e. 10 ppm, growth tended to be promoted, whereas at 100 ppm growth appeared to be suppressed. Continued testing of the bioregulator's effect on growth and physiology of other tree species is recommended.     

Identification of Cylindrocladium sp. causing damping-off disease of Japanese black pine (Pinus thunbergii) and factors affecting the disease severity in a black locust (Robinia pseudoacacia)-dominated area

The regeneration of Japanese black pine (Pinus thunbergii) seedlings is inhibited in a black locust (Robinia pseudoacacia)-dominated area. We examined the presence of pathogenic fungi in Japanese black pine seedlings in the area in order to determine the effect of pathogenic fungi on the inhibition of regeneration. When Japanese black pine seedlings were planted in the soil obtained from a black locust-dominated area, all of the seedlings died under low-intensity light conditions, whereas 84% of the seedlings survived in the soil obtained from a Japanese black pine-dominated area under the same light conditions. One fungus was isolated from 48.7% of the dead pine seedlings and was identified as Cylindrocladium pacificum Kang, Crous & Schoch, based on the morphological characteristics, growth, and DNA analysis. This fungus was also isolated from 50% of the dead pine seedlings in 2005 and 66.7% of the seedlings in 2006—both were planted in a black locust-dominated area. The virulence of this fungus increased under high-nitrogen and/or low-intensity light conditions. These results reveal the possibility that the soil eutrophication and shading by the black locust are conducive to a severe damping-off disease and threaten the survival and regeneration of Japanese black pine seedlings.     

Field survival of containerized red and jack pine seedlings inoculated with mycelial slurries of ectomycorrhizal fungi

Red pine and jack pine seedlings growing in styroblocks were inoculated 8 wk after sowing with mycelium/agar slurries of 3 mycorrhizal fungi (Laccaria bicolor, Scleroderma citrinum, and an unidentified basidiomycete), and one suspected mycorrhizal fungus (Cantharellula umbonata). Seedlings inoculated with L. bicolor developed mycorrhizae earlier and in greater numbers than the other inoculation treatments, with red pine out-performing jack pine in both respects. At 34 wk following sowing, seedlings were outplanted on a cleared xeric site in Baraga Co., in Michigan's Upper Peninsula. Seedlings inoculated with C. umbonata failed to form mycorrhizae and were not outplanted. Inoculation treatments did not affect shoot or root weight at outplanting. Red pine inoculated with L. bicolor averaged 21% and 19% greater survival compared with control seedlings after one and two years in the field, respectively. Other inoculation treatments failed to increase seedling survival for either tree species. Jack pine demonstrated higher overall survival than did red pine for both years in all corresponding inoculation treatments.     

Gas exchange,water relations and morphology of yellow-cedar seedlings and stecklings before planting and during field establishment

Information is lacking on how yellow-cedar (Chamaecyparis nootkatensis (D. Don.) Spach) stecklings compare with seedlings in their response to potentially limiting field site environmental conditions. Before planting, yellow-cedar seedlings and stecklings were characterized at 5 and 22°C root temperatures for gas exchange parameters (n=6), root growth capacity (n=12) and plant water movement (n=6) using a fully randomized design in a controlled environment growth room. Seedlings, compared with stecklings (t-test), showed a faster (p=0.01) recovery of net photosynthesis (P_n) and stomatal conductance (g_wv) after root temperature increased from 5 to 22°C. Seedlings had greater (p=0.05) root growth capacity (number of new roots 0.5 cm in length) than stecklings after 14 days at 22°C and after 14 days at 5°C followed by 15 days at 22°C. Seedlings had a lower (p=0.01) resistance to plant water movement measured after 29 days (14 days at 5°C followed by 15 days at 22°C root temperature).Seedlings and stecklings were planted on a coastal British Columbia field site and monitored for 1) morphological development and physiological response to ambient site conditions throughout the first growing season, and 2) physiological response to drought conditions during late summer. Both trials used a fully randomized design and stock type means were compared using t-tests (p=0.10, 0.05, and 0.01). Initially, seedlings were taller (p=0.01) than stecklings but otherwise morphologically similar (n=20). Over a 35 week period after planting, stecklings had greater incremental height growth, while seedlings produced a greater (p=0.01) number of roots extending out of the root plug. Seedlings and stecklings (n=6) had similar P_n and g_wv patterns under late spring environmental conditions. However, seedlings had greater P_n and g_wv under cold temperature (p=0.05 and p=0.1, respectively) and drought conditions (p=0.05 and p=0.01, respectively), but during conditions of high evaporative demand and adequate soil moisture, stecklings had greater (p=0.01) P_n and g_wv. Seedlings had higher (p=0.01) daily integrated shoot water potential values (D) at different times during the growing season, and maintained higher (p=0.05) predawn (_pd) shoot water-potential values, higher (D) (p=0.01) and lower (p=0.01) resistance to plant water movement (R_SPAC) during drought conditions.     

Relationship between the incidence of pine wilt disease and the drainage area

We examined the relationship between the incidence of pine wilt disease and moisture conditions in the stand level ofPinus densiflora Sieb. et Zucc. forests in the warm-temperate zone of the western part of Japan. For this analysis, pine trees killed by pine wilt disease were distinguished from the suppressed trees by their position in the layer of the pine forest stand. The drainage area, which is small in the upper part and large in the lower part of the slope, was adopted for representing the moisture conditions in the soil of the slope. The percentage of the pine trees killed by pine wilt disease increased as the size of the drainage area increased. This result suggested that the incidence of pine wilt disease tended to be high in areas with moist conditions. Pine trees attacked by the pinewood nematode die from extensive water deficit due to tracheid cavitations. Pine wilt disease mainly emerges in the summer when the soil water conditions become especially severe, and the radical water stress is thought to accelerate the disease. It was assumed that pine trees in the plots with the small drainage area resisted the influence of the attack of the pinewood nematode because pine trees in the plots with the small drainage area encountered long-term water stress and acquired drought tolerance. Pine trees in the plots with the large drainage area were presumed to be well established in the moist conditions and not to have acquired drought tolerance. The drought tolerance of pine trees was thought to be an important factor in resistance to pine wilt disease.     

The penetration rate of laser pulses transmitted from a small-footprint airborne LiDAR: a case study in closed canopy, middle-aged pure sugi (Cryptomeria japonica D. Don) and hinoki cypress (Chamaecyparis obtusa Sieb. et Zucc.) stands in Japan

Penetration rates of laser pulses transmitted from a small-footprint airborne light detection and ranging (LiDAR) system in closed-canopy, middle-aged (40–50 years old) hinoki cypress (Chamaecyparis obtusa Sieb. et Zucc.) and sugi (Cryptomeria japonica D. Don) plantations that had similar levels of canopy openness were studied. The number of transmitted pulses was 107 427 points/ha for the hinoki cypress stand and 122 883 points/ha for the sugi stand. The penetration rates of the first (P_f) and second pulses (P_s) that reached the ground were 0.1% and 1.0% in the hinoki cypress stand, and 2.6% and 5.5% in the sugi stand, respectively. Penetration rates for the total number of pulses (P_f+s) that reached the ground within each stand were 1.1% and 8.1%, respectively, and values for P_f+s, P_f, and P_s for sugi and hinoki cypress stands were significantly different (P < 0.001). The results of this study indicated that the generation of accurate digital terrain models in dense hinoki cypress stands with complex topographies using only the data from laser pulses with poor laser penetration rates is likely to be difficult.     

Grass interference limits resource availability and reduces growth of juvenile red pine in the field

The effects of competing grasses on resource availability, growth and ecophysiological characteristics of 3-0 red pine (Pinus resinosa Ait.) seedlings were examined the first two years following outplanting in Anoka County, Minnesota, USA. Equal numbers of seedlings were planted into suppressed and undisturbed grass communities in a sandy soil. Grass suppression was maintained throughout the first growing season, but partially discontinued thereafter on the site. During the first field season interference from grass reduced pine seedling root collar diameter, needle length, number of new root tips, and lateral root length by over 40%. Mean pre-dawn needle water potential was 0.55 MPa lower in undisturbed grass plots during a brief drought in year one, but otherwise water stress was not significantly (p=0.05) influenced by grass interference. The presence of grass also reduced, up to 50%, the photosynthetically active radiation reaching the seedling canopy. At the end of year one, total biomass N, P, K, and Ca content were significantly (p=0.05) less in seedlings growing in the undisturbed grass community. Nitrogen was deficient in seedlings growing in grass. After two growing seasons, seedling shoot length (p=0.03), root collar diameter (p=0.001), and needle length (p=0.001) were significantly less (40, 54 and 20%, respectively) for seedlings growing in undisturbed grass. Seedling growth reductions induced by grass competition were associated with multiple environmental stressors in the field and not restricted to water stress as was observed in earlier studies with pine species at low and mid-latitude sites.     

Prescribed burning and hexazinone herbicide as release treatments in a sapling hardwood-loblolly pine stand

To compare release treatments, a randomized complete block study was established in a 7-year-old hardwood-loblolly pine (Pinus taeda L.) stand in central Louisiana established using chopping and burning. There were 5 blocks of 3 treatments each: (1) check, (2) hexazinone applied once, and (3) prescribed backfiring applied twice. The first burn in December 1985 (7 years after site preparation) had a fire intensity of 90 kJ/s/m. The hexazinone herbicide was applied in April 1986 (the 8th year after site preparation) with a metered spotgun applicator at a rate of 3.0 kg active ingredient/ha. The second burn in March 1989 (the 11th year after site preparation) had a fire intensity of 106 kJ/s/m.The two prescribed burns increased the number of stems less than 1.5 m tall from 1,380 to 2,960/ha red maple (Acer rubrum L.), blackgum (Nyssa sylvatica var. bifora), and sweetgum (Liquidambar styraciflua L.). Burning reduced the number of loblolly pines less than 2.0 m tall, which resulted in a significant increase in average loblolly pine height and diameter. Loblolly pine comprised 62 and 59% of the basal area on the check and burn treatments, respectively, 11 years after site preparation. Hexazinone reduced the number of blackgum, sweetgum, and oak (Quercus spp.) from 6,100 to 4,560 stems/ha and resulted in significantly less hardwood tree basal area than found on the check or burn treatments. Therefore, the herbicide treatment resulted in principally a loblolly pine stand (over 80% of the total tree basal area was pine) four years after hexazinone application.     

Allometric model of the Reineke equation for Japanese cypress (<Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis>) and red pine (<Emphasis Type="Italic">Pinus densiflora</Emphasis>) stands

An allometric model that explains the mechanism of the difference in the slope of the Reineke equation (A) among species was proposed based on the allometric relationships of mean tree height (H) to quadratic mean diameter D (H ∝ D ^θ ) and stand density N (H ∝ N ^δ ), i.e., A = θ/δ. The proposed model was fitted to Japanese cypress (Chamaecyparis obtusa Endl.) and red pine (Pinus densiflora) stands. The allometric exponents θ and δ were, respectively, 0.8995 and −0.5000 for cypress and 0.8612 and −0.6619 for pine. The difference between cypress and pine was significant for δ but not for θ. Inserting the exponents into the model resulted in predicted slopes of −1.7991 for cypress and −1.3011 for pine. The difference in the slope of the Reineke equation between the two species was produced by characteristics related to the tree crown, rather than characteristics related to stem slenderness. The proposed model enables us to estimate the slope of the Reineke equation from commonly measured stand attributes, such as mean tree height and quadratic mean diameter. Therefore, the proposed model is expected to be practical and convenient for estimating the slope of the Reineke equation and for explaining the mechanism of its variation among species. The model should be also accepted as a generalized model of the stand density versus quadratic mean diameter relationship, whereas the original Reineke equation should be seen as a specific case of this model.