Dead Wood is Buried and Preserved in a Labrador Boreal Forest

Large amounts (36.4 Mg ha⁻¹ or 179 m³ ha⁻¹) of buried dead wood were found in overmature (146–204-year-old) black spruce (Picea mariana (Mill.) B.S.P.) forests in the high boreal region of eastern Canada. Amounts of this size indicate that burial reduces rates of wood decay producing an important component of long-term carbon (C) storage. Radiocarbon-derived ages of black spruce stems buried near the bottom of the organic soil horizon at three old-growth sites were up to 515 years old. Together with information on current stand age, this suggests that the stems have been dead for more than 250 years. Most aboveground dead wood decays or becomes fragmented within about 70 years of tree death in these forests. The presence of old yet well-preserved buried wood suggests that decay rates are greatly reduced when downed dead wood is quickly overgrown by moss. Thus, the nature and type of ground-layer vegetation influences the accumulation of organic matter in these forests. This process of dead wood burial and the resultant addition to a large and long-enduring belowground C pool should be considered when estimating dead wood abundance for habitat or forest C accounting and cycling.     

Forests regenerating after clear-cutting function as habitat for bryophyte and lichen species of conservation concern

The majority of managed forests in Fennoscandia are younger than 70 years old but yet little is known about their potential to host rare and threatened species. In this study, we examined red-listed bryophytes and lichens in 19 young stands originating from clear-cutting (30-70 years old) in the boreal region, finding 19 red-listed species (six bryophytes and 13 lichens). We used adjoining old stands, which most likely never had been clear-cut, as reference. The old stands contained significantly more species, but when taking the amount of biological legacies (i.e., remaining deciduous trees and dead wood) from the previous forest generation into account, bryophyte species number did not differ between old and young stands, and lichen number was even higher in young stands. No dispersal effect could be detected from the old to the young stands. The amount of wetlands in the surroundings was important for bryophytes, as was the area of old forest for both lichens and bryophytes. A cardinal position of young stands to the north of old stands was beneficial to red-listed bryophytes as well as lichens. We conclude that young forest plantations may function as habitat for red-listed species, but that this depends on presence of structures from the previous forest generation, and also on qualities in the surrounding landscape. Nevertheless, at repeated clear-cuttings, a successive decrease in species populations in young production stands is likely, due to increased fragmentation and reduced substrate amounts. Retention of dead wood and deciduous trees might be efficient conservation measures. Although priority needs to be given to preservation of remnant old-growth forests, we argue that young forests rich in biological legacies and located in landscapes with high amounts of old forests may have a conservation value.     

Bryophyte species richness and composition in young forests regenerated after clear-cut logging versus after wildfire and spruce budworm outbreak

The disturbance regime in mixed-wood forests of eastern Canada is characterized by both natural disturbances including wildfires and insect outbreaks as well as forestry. The understanding of how understorey plant assemblages respond to different disturbances is mostly limited to short-term wildfire-logging comparisons of vascular plants. Here, we compare patterns of species richness and composition of four bryophyte guilds in young forests (approx. 40 years old) regenerating after clear-cut logging, wildfire, and spruce budworm outbreak. In addition, young forests were compared with mature spruce-fir dominated stands (approx. 90 years old). Although similar in overall species richness at the scale of 1,000 m² all young forest types were compositionally distinct with fewer species than mature forests. Stands developed after spruce budworm outbreaks had the highest canopy cover values and the highest surface area of coarse woody debris. These stands had similar numbers of woody debris species as mature forests and were closest to mature forests in species composition. Wildfire-disturbed sites were dominated by deciduous trees and a high number of treebase species. Finally, young managed forest had the highest number of forest floor bryophytes at the scale of 100 m² among the three young forest types, but was compositionally far from mature forests in their woody debris flora. In conclusion, young forests regenerating after natural disturbances are distinctly different from young forests regenerated after clear-cutting and if natural disturbances are eliminated certain species (e.g., epixylic and treebase species) might become more restricted to older stands in the landscape.     

The herb and dwarf shrubs colonization of decaying logs in subalpine forest in the Polish Tatra Mountains

This is a study of the colonization pattern of herbs and dwarf shrubs on rotten logs in subalpine spruce forests (Plagiothecio Piceetum) in the Tatra Mountains. On four study plots (total area 1.43 ha.) all dead logs were measured and the decomposition stage was estimated using the 8-degree scale. For each log the cover of all vascular species, bryophytes and lichens was determined according to the methods of classical phytosociology. Constancy and an index of coverage were calculated for all vascular species growing on logs. The total volume of logs was relatively high (93 m³ ha^–1) and constituted 22% of the volume of living trees. Logs and stumps covered 411 m² ha^–1. These values are similar to those known from natural spruce forest from Carpathians and Scandinavia. The 8 stages of decomposition were equally represented, which indicates a constant supply of dead wood to the forest floor over time. The colonization of dead wood starts with lichens, followed by bryophytes and finally herbs and tree saplings. The first vascular plant colonists of dead logs appear at decay stage nr. 3 at least 20 years after tree death. The most suitable condition for most of the herb species corresponds to decay stage nr. 6 ca. 50 years after tree death. The herb cover is distinctively dominated by Vaccinium myrtillus. Simultaneously with herb species, tree seedlings colonize the logs. Constancy and abundance of Norway spruce saplings increases with advanced decomposition. It seems that the herb cover of logs does not hinder the regeneration of spruce.     

The effect of different anthropogenic disturbances on litterfall of a dominant pioneer rain forest tree in Gabon

Tropical rain forests worldwide are affected by anthropogenic disturbances, and secondary forests that develop afterwards are often dominated by pioneer tree species, but the consequences of different anthropogenic disturbances on nutrient cycling are poorly understood. Because most nutrient cycling in tropical rain forests occurs through litterfall and in the soil organic layer, we measured litterfall of a widespread and dominant pioneer tree, okoume (Aucoumea klaineana, Burseraceae) in Gabon, in one forest previously disturbed by logging and in another by agriculture. Litterfall of okoume trees, measured over 14 months, was 18.2 Mg ha⁻¹ year⁻¹ in the formerly logged forest, which was 72.6% greater than in the forest more recently disturbed by agriculture (10.6 Mg ha⁻¹ year⁻¹). Litter decomposition rates were more rapid in the formerly logged forest, and this may explain why litter thickness was 32% lower in the formerly logged forest, despite the greater litterfall within it. Our results highlight that two widespread anthropogenic disturbances in Gabonese rain forests have significantly different consequences on litterfall of a dominant tree and therefore are likely to have different effects on nutrient cycling and forest ecosystem function.     

Effects of experimental stem burial on radial growth and wood anatomy of pedunculate oak

In dendrogeomorphology, abrupt changes in wood anatomy are frequently used to date the exact year of burial and exposure events. However, few studies have addressed the precision and underlying mechanisms of these changes. In a field experiment, performed in a drift-sand area in the Netherlands, we buried the stems of mature pedunculate oak trees (Quercus robur L.) up to a height of 50 cm and analysed the responses in ring width and earlywood-vessel characteristics, while monitoring the course of temperature above and below the soil surface.After 3 years of stem burial, we found no significant differences in ring width and earlywood-vessel characteristics between control and buried trees both above and below the burial level. Burial however strongly reduced temperature amplitude and the occurrence of sub-zero temperatures around the buried stems. All buried trees formed epitropic roots that grew upward into the new sediment layer, but no adventitious roots were formed on the buried stems. Irrespective of the burial treatments, we found that the mean ring width was largest at the original stem base and lowest at breast height. In contrast, vessel sizes were significantly larger at breast height compared with the stem base. Differences in vessel density barely differed between years and heights.In our field experiment on mature pedunculate oak trees, the burial of stems by 50 cm of drift sand did not induce any local growth suppression or detectable changes in wood anatomy. As wood-anatomical changes in response to burial have previously been reported for trees that had formed adventitious roots, we stress the role of adventitious-root formation as a possible trigger behind the local changes in wood anatomy, reflecting a functional change of a buried stem towards a root. Based on our field experiment, it seems unlikely that years of shallow or moderate burial events (≤50 cm) can be reconstructed using the wood structure of buried stems. As epitropic roots develop quickly after burial, dating such roots may potentially yield better estimates of burial events. Further research on the relation between adventitious root and changes in stem anatomy is needed to ascertain the precision of dating sand-burial events using tree rings.     

Diversity of saproxylic bryophytes in old‐growth and managed beech forests in the central Balkans

Abstract

The diversity of saproxylic bryophyte species in beech forest stands from the wide region of the central Balkans (i.e. Serbia and Montenegro) was studied, and this study is the first of such a type in SE Europe. Comparison of preserved old‐growth and managed forests were made. Bryophyte species diversity is higher in primeval forest stands where the spectra of dead wood in various decaying stages of its dynamics are present. The ecological group of epixylic specialists is predominant among the bryophytes recorded. Threatened bryophyte species occur in old‐growth beech stands. The dead wood as habitat together with some other factors are extremely important for the surviving of epixylic bryophyte; so these species can be used as bioindicator bryophyte species of old‐growth or managed and structured forest ecosystems.     

Implications of floristic and environmental variation for carbon cycle dynamics in boreal forest ecosystems of central Canada

Abstract. Species composition, detritus, and soil data from 97 boreal forest stands along a transect in central Canada were analysed using Correspondence Analysis to determine the dominant environmental/site variables that differentiate these forest stands. Picea mariana stands were densely clustered together on the understorey DCA plot, suggesting a consistent understorey species composition (feather mosses and Ericaceae), whereas Populus tremuloides stands had the most diverse understorey species composition (ca. 30 species, mostly shrubs and herbs). Pinus banksiana stands had several characteristic species of reindeer lichens (Cladina spp.), but saplings and Pinus seedlings were rare. Although climatic variables showed large variation along the transect, the CCA results indicated that site conditions are more important in determining species composition and differentiating the stand types. Forest floor characteristics (litter and humus layer, woody debris, and drainage) appear to be among the most important site variables. Stands of Picea had significantly higher average carbon (C) densities in the combined litter and humus layer (43530 kg‐C.ha^‐1) than either Populus (25 500 kg‐C.ha^‐1) or Pinus (19 400 kg‐C.ha^‐1). The thick surface organic layer in lowland Picea stands plays an important role in regulating soil temperature and moisture, and organic‐matter decomposition, which in turn affect the ecosystem C‐dynamics. During forest succession after a stand‐replacing disturbance (e.g. fires), tree biomass and surface organic layer thickness increase in all stand types as forests recover; however, woody biomass detritus first decreases and then increases after ca. 80 yr. Soil C densities show slight decrease with ages in Populus stands, but increase in other stand types. These results indicate the complex C‐transfer processes among different components (tree biomass, detritus, forest floor, and soil) of boreal ecosystems at various stages of succession.     

Effects of harvesting on spatial and temporal diversity of carbon stocks in a boreal forest landscape

Carbon stocks in managed forests of Ontario, Canada, and in harvested wood products originated from these forests were estimated for 2010–2100. Simulations included four future forest harvesting scenarios based on historical harvesting levels (low, average, high, and maximum available) and a no‐harvest scenario. In four harvesting scenarios, forest carbon stocks in Ontario's managed forest were estimated to range from 6202 to 6227 Mt C (millions of tons of carbon) in 2010, and from 6121 to 6428 Mt C by 2100. Inclusion of carbon stored in harvested wood products in use and in landfills changed the projected range in 2100 to 6710–6742 Mt C. For the no‐harvest scenario, forest carbon stocks were projected to change from 6246 Mt C in 2010 to 6680 Mt C in 2100. Spatial variation in projected forest carbon stocks was strongly related to changes in forest age (r = 0.603), but had weak correlation with harvesting rates. For all managed forests in Ontario combined, projected carbon stocks in combined forest and harvested wood products converged to within 2% difference by 2100. The results suggest that harvesting in the boreal forest, if applied within limits of sustainable forest management, will eventually have a relatively small effect on long‐term combined forest and wood products carbon stocks. However, there was a large time lag to approach carbon equality, with more than 90 years with a net reduction in stored carbon in harvested forests plus wood products compared to nonharvested boreal forest which also has low rates of natural disturbance. The eventual near equivalency of carbon stocks in nonharvested forest and forest that is harvested and protected from natural disturbance reflects both the accumulation of carbon in harvested wood products and the relatively young age at which boreal forest stands undergo natural succession in the absence of disturbance.     

Relationship between decomposition of cellulose in the soil and tree stand characteristics in natural boreal forests

The relationship between the decomposition of cellulose placed on and buried in the forest floor and various tree stand characteristics was studied at sites with minimal anthropogenic influence. The 22 study sites, including both forested upland and peatland plots, were clustered in 4 catchments between 61°–69° N in Finland. The stands were 60 to 320 years old and composed of varying proportions of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies Karst.) and deciduous species (mainly t Betula spp.). Cellulose strips (softwood pulp) were placed on the forest floor surface and buried at four depths down to 5 cm for two 1-year periods and the weight loss measured. Decomposition did not significantly correlate with stand age, but was strongly and positively correlated with stand basal area, mean tree height and stem volume. This was valid at all depths, and even after differences due to climate between catchments had been taken into account. The stem volume of Scots pine dominated plots had the highest correlation. Our results showed that decomposition of organic matter on and in the forest floor is related to the stand characteristics. This relationship should be considered when comparing soil processes different stands, even when comparing stands of the same tree species composition.     

Tree mycorrhizal type predicts within‐site variability in the storage and distribution of soil organic matter

Forest soils store large amounts of carbon (C) and nitrogen (N), yet how predicted shifts in forest composition will impact long‐term C and N persistence remains poorly understood. A recent hypothesis predicts that soils under trees associated with arbuscular mycorrhizas (AM) store less C than soils dominated by trees associated with ectomycorrhizas (ECM), due to slower decomposition in ECM‐dominated forests. However, an incipient hypothesis predicts that systems with rapid decomposition—e.g. most AM‐dominated forests—enhance soil organic matter (SOM) stabilization by accelerating the production of microbial residues. To address these contrasting predictions, we quantified soil C and N to 1 m depth across gradients of ECM‐dominance in three temperate forests. By focusing on sites where AM‐ and ECM‐plants co‐occur, our analysis controls for climatic factors that covary with mycorrhizal dominance across broad scales. We found that while ECM stands contain more SOM in topsoil, AM stands contain more SOM when subsoil to 1 m depth is included. Biomarkers and soil fractionations reveal that these patterns are driven by an accumulation of microbial residues in AM‐dominated soils. Collectively, our results support emerging theory on SOM formation, demonstrate the importance of subsurface soils in mediating plant effects on soil C and N, and indicate that shifts in the mycorrhizal composition of temperate forests may alter the stabilization of SOM.     

Size and Structure of Fine Root Systems in Old-growth and Secondary Tropical Montane Forests (Costa Rica)

The fine root systems of three tropical montane forests differing in age and history were investigated in the Cordillera Talamanca, Costa Rica. We analyzed abundance, vertical distribution, and morphology of fine roots in an early successional forest (10–15 years old, ESF), a mid‐successional forest (40 years old, MSP), and a nearby undisturbed old‐growth forest (OGF), and related the root data to soil morphological and chemical parameters. The OGF stand contained a 19 cm deep organic layer on the forest floor (i.e., 530 mol C/m²), which was two and five times thicker than that of the MSF (10 cm) and ESF stands (4 cm), respectively. There was a corresponding decrease in fine root biomass in this horizon from 1128 g dry matter/m² in the old‐growth forest to 337 (MSF) and 31 g/m² (ESF) in the secondary forests, although the stands had similar leaf areas. The organic layer was a preferred substrate for fine root growth in the old‐growth forest as indicated by more than four times higher fine root densities (root mass per soil volume) than in the mineral topsoil (0–10 cm); in the two secondary forests, root densities in the organic layer were equal to or lower than in the mineral soil. Specific fine root surface areas and specific root tip abundance (tips per unit root dry mass) were significantly greater in the roots of the ESF than the MSF and OGF stands. Most roots of the ESF trees (8 abundant species) were infected by VA mycorrhizal fungi; ectomycorrhizal species (Quercus copeyemis and Q. costaricensis) were dominant in the MSF and OGF stands. Replacement of tropical montane oak forest by secondary forest in Costa Rica has resulted in (1) a large reduction of tree fine root biomass; (2) a substantial decrease in depth of the organic layer (and thus in preferred rooting space); and (3) a great loss of soil carbon and nutrients. Whether old–growth Quercus forests maintain a very high fine root biomass because their ectomycorrhizal rootlets are less effective in nutrient absorption than those of VA mycorrhizal secondary forests, or if their nutrient demand is much higher than that of secondary forests (despite a similar leaf area and leaf mass production), remains unclear.     

Interactions with successional stage and nutrient status determines the life‐form‐specific effects of increased soil temperature on boreal forest floor vegetation

The boreal forest is one of the largest terrestrial biomes and plays a key role for the global carbon balance and climate. The forest floor vegetation has a strong influence on the carbon and nitrogen cycles of the forests and is sensitive to changes in temperature conditions and nutrient availability. Additionally, the effects of climate warming on forest floor vegetation have been suggested to be moderated by the tree layer. Data on the effects of soil warming on forest floor vegetation from the boreal forest are, however, very scarce. We studied the effects on the forest floor vegetation in a long‐term (18 years) soil warming and fertilization experiment in a Norway spruce stand in northern Sweden. During the first 9 years, warming favored early successional species such as grasses and forbs at the expense of dwarf shrubs and bryophytes in unfertilized stands, while the effects were smaller after fertilization. Hence, warming led to significant changes in species composition and an increase in species richness in the open canopy nutrient limited forest. After another 9 years of warming and increasing tree canopy closure, most of the initial effects had ceased, indicating an interaction between forest succession and warming. The only remaining effect of warming was on the abundance of bryophytes, which contrary to the initial phase was strongly favored by warming. We propose that the suggested moderating effects of the tree layer are specific to plant life‐form and conclude that the successional phase of the forest may have a considerable impact on the effects of climate change on forest floor vegetation and its feedback effects on the carbon and nitrogen cycles, and thus on the climate.     

Role of the Siberian flying squirrel as an umbrella species for biodiversity in northern boreal forests

One of the potentially useful indirect shortcut methods in biodiversity conservation is the umbrella species concept. An umbrella species can be seen relatively demanding for the size of the area and probably also for certain habitat types: conservation management for the umbrella species would thus encompass other species preferring similar habitats but with smaller area requirements. As such, it has a comprehensive spatial aspect for landscape planning. We tested the role of the Siberian flying squirrel (Pteromys volans) as an umbrella species for wood dependent species among red-listed and old-growth forest associated polypores, epiphytic lichens and beetles. Flying squirrels inhabit home ranges of several to tens of hectares, and prefer mature spruce-dominated (Picea abies) mixed forests, which often have high amounts of dead wood. We carried out species surveys and trappings during 1 year from 20 mature spruce-dominated forest stands (altogether 162 ha), of which 12 were occupied by the flying squirrel. The amount of dead wood was higher in occupied stands than in unoccupied stands. We also found a tendency for a higher number of species and number of records in occupied stands, a relationship mostly due to the polypore species. The presence of the flying squirrel may reflect the habitat availability for species depending on dead and living wood, and assist in site selection of conservation areas. We suggest that the flying squirrel has potential as an umbrella species to partly enhance maintenance of biodiversity in northern boreal forests in Finland.     

华北地区针叶林下凋落物层化学性质的研究

森林凋落物中贮积了大量的有机、无机养分物质,它是森林土壤自然肥力的重要来源之一。在森林生态系统中,养分物质的内部循环主要是通过凋落物来实现的。在分解、转化过     

Forest history and the development of old‐growth characteristics in fragmented boreal forests

Questions: Can small and isolated high‐conservation value forests (e.g. designated woodland key habitats) maintain old‐growth forest characteristics and functionality in fragmented landscapes? To what extent have past disturbances (natural and anthropogenic) influenced the development of old‐growth characteristics of these forests? How long does it take for selectively cut stands to attain conditions resembling old‐growth forests? Location: Southern boreal zone of central Sweden. Methods: We linked multiple lines of evidence from historical records, biological archives, and analyses of current forest structure to reconstruct the forest history of a boreal landscape, with special emphasis on six remaining core localities of high‐conservation value forest stands. Results: Our reconstructions revealed that several of these stands experienced wildfires up to the 1890s; all had been selectively harvested in the late 1800s; and all underwent substantial structural and compositional reorganization over the following 100‐150 years. This time interval was sufficient to recover considerable amounts of standing and downed dead wood (mean 60.3 m³ ha^?1), a range of tree ages and sizes (mean basal area 32.6 m² ha^?1), and dominance of shade‐tolerant spruce. It was insufficient to obtain clearly uneven tree age structures and large (>45 cm diameter) living and dead trees. Thus, these forests contain some, but not all, important compositional and structural attributes of old‐growth forests, their abundance being dependent on the timing and magnitude of past natural and anthropogenic disturbances. Our landscape‐level analysis showed marked compositional and structural differences between the historical forest landscape and the present landscape, with the latter having a greater proportion of young forests, introduction of non‐native species, and lack of large trees and dead wood. Conclusions: The remnant high‐conservation value stands were not true representatives of the pre‐industrial forests, but represent the last vestige of forests that have regenerated naturally and maintained a continuous tree cover. These traits, coupled with their capacity for old‐growth recovery, make them valuable focal areas for conservation.     

Experimental transplants reveal strong environmental effects on the growth of non‐vascular epiphytes in Afromontane forests

Transplant studies can provide valuable information on the growth responses of epiphytic bryophytes and lichens to environmental factors. We studied the growth of six epiphyte species at three sites in moist Afromontane forests of Taita Hills, Kenya. With 558 pendant transplants, we documented the growth of four bryophytes and two lichens over 1 yr. The transplants were placed into the lower canopy of one forest site in an upper montane zone, and two forest sites in a lower montane zone. Several pendant moss species grew very well in the cool and humid environment of the upper montane forest, with some transplants more than doubling their biomass during the year. Conversely, all transplanted taxa performed poorly in the lower montane zone, presumably because of the unfavorable combination of ample moisture with excessive warmth and insufficient light which characterizes the lower canopy in dense lower montane forests. The results demonstrate that pendant transplants can be used for monitoring growth of non‐vascular epiphytes in tropical forests. The starting weight of 0.25 g for pendant transplants worked well and can be recommended for future studies.     

The soil organic carbon in particle-size separates under different regrowth forest stands of north eastern Costa Rica

Despite the importance of the secondary forest (SF) in tropical areas, few studies have quantified the soil organic carbon (SOC) pool in Costa Rica. Most of the studies conducted to date in this country have focused mainly on changes in the soil C pool following conversion of forests to pastures, which is the predominant land use in the tropics. The aim of this study was to measure SOC concentration and pool in particle-size fractions down to 50 cm depth in four SF stands regenerating from different intensities of prior land use in loamy sand and sandy loam soils of northeast Costa Rica: (i) a gallery forest (GF), (ii) a 15-year-old SF enriched with commercially planted native trees (15SF), (iii) a 25-year-old SF (25SF), and (iv) an abandoned Theobromma cacao plantation >60 years old (60SF). Additional objectives were (1) to determine the relationship of SOC concentration with selected physical and chemical soil properties, and (2) to establish the key determinants of the depth distribution of SOC in order to identify meaningful trends in the SOC pool. The SOC pool was highest under the 60SF (221.4 Mg C ha⁻¹) followed by the 15SF (212.1 Mg C ha⁻¹), the 25SF (195.9 Mg C ha⁻¹) and the lowest in the GF (183.5 Mg C ha⁻¹). The SOC concentration decreased significantly from 59.7 to 94.1 g kg⁻¹ in the 0–10 cm layer down to 31.0 to 45.5 g kg⁻¹ in the 40–50 cm layer in all forest stands. The fine silt + clay fraction contained the highest values of SOC concentration in all forest stands. Soil texture and the age of the SF were identified as the main factors that explained the variability in SOC. The age of SF stand influenced the distribution of size class aggregates and SOC.     

甘肃小陇山不同针叶林凋落物量、养分储量及持水特性

通过野外实地观测和室内分析相结合的方法对甘肃小陇山地区6种主要针叶林凋落物层的蓄积量、持水特性及养分储量进行了调查,结果表明:云杉林凋落物层的蓄积量最大,松林次之,落叶松林最小。同一林分林下半分解层凋落物的持水能力比分解层强;在不同林分类型中,凋落物层的最大持水率、最大拦蓄率和有效拦蓄率的大小顺序为:落叶松林>松林>云杉林,而最大拦蓄量和有效拦蓄量的大小顺序为:云杉林>松林>落叶松林。同一林分类型中,分解层凋落物中养分储量最多,未分解层最少;不同林分类型中各个营养元素的储量均表现出:云杉林>松林>落叶松林。凋落物蓄积量与营养元素储量密切相关,最大持水量与凋落物储量和凋落物层各元素储量之间呈显著的正相关性。