A comparison of the reproductive parameters of females of two reptile tick species

In comparisons of females of two reptile tick species Aponomma hydrosauri and Amblyomma limbatum, Ap. hydrosauri was initially larger, and after mating on the host engorged faster and remained attached for a shorter time before completing engorgement and detaching. Amb. limbatum had a longer period of engorgement, and achieved a greater engorged weight. Engorged Amb. limbatum females laid significantly more eggs than equivalent sized Ap. hydrosauri. Although the two species are ecologically similar and were collected from the same site for this study, their reproductive differences probably reflect adaptations to different conditions in their largely allopatric ranges.     

Delayed mating and the reproductive fitness of aponomma hydrosauri (Acari: Ixodidae)

, and 1992. Delayed mating and the reproductive fitness of Aponomma hydrosauri (Acari: Ixodidae). International Journal for Parasitology 22: 1197–1200. This study examines whether delayed mating influences the reproductive fitness of female reptile ticks, Aponomma hydrosauri. Delayed mating was induced by the prevention of male attachment to hosts for 20 or 40 days after females had been attached. The results showed that delayed mating had no significant influence on the number of viable progeny produced by female ticks. This may represent an important advantage for colonizing Ap. hydrosauri females in marginal population areas, particularly at parapatric boundaries.     

格氏栲幼苗叶绿素荧光和非结构性碳水化合物积累对种子散布位置的响应

植物种子从母树掉落形成土壤种子库时,凋落物或土壤是其最初接触的物理环境,种子所处位置(种子在凋落物上层、土壤表层或凋落物下层)影响了幼苗天然更新进程。模拟格氏栲种子在凋落物上层(种子下层铺垫2和4 cm凋落物)、土壤表层(无凋落物)及凋落物下层(种子上层覆盖2、4、6和8 cm凋落物)等3种不同散布位置,探讨种子散布位置对幼苗叶绿素荧光特性、非结构性碳水化合物、比叶面积、叶干物质含量和养分含量的影响。结果表明:不同散布位置的幼苗单位面积的叶氮含量与可溶性糖、非结构性碳水化合物含量呈显著正相关,与比叶面积呈显著负相关。适宜凋落物覆盖(2和4 cm)的幼苗通过提高叶绿素相对含量、可溶性糖含量、非结构性碳水化合物含量、叶干物质含量和单位面积的叶氮含量和叶磷含量,降低比叶面积等的资源获取策略来实现自身快速生长需求。无凋落物和深层凋落物覆盖(6和8 cm)的幼苗采取高单位重量的叶氮含量和比叶面积,低叶干物质含量和非结构性碳水化合物含量的资源保守型策略以截获更多有效光资源,进而弥补深层凋落物带来的郁闭环境,降低幼苗因“碳饥饿”而死亡的几率。下层铺垫凋落物的幼苗通过在叶片储藏淀粉,降低叶片光合组织消耗能量(低PSⅡ最大光化学效率)等维持幼苗生长。熵值法综合分析表明,浅层凋落物覆盖(2 cm)对格氏栲幼苗生长的促进作用最为显著,未来可通过调节天然林凋落物层厚度以促进格氏栲幼苗生长与更新。     

The salivary glands of Hyalomma anatolicum anatolicum: structural changes during attachment and feeding

The histology and ultrastructure of the salivary glands of male and female H.a. anatolicum ticks have been examined m unfed and feeding ticks with special emphasis on aspects related to the feeding process. The salivary glands of H.a. anatolicum consisted of three types of acinus (acinus I, II and III) in females and an additional type IV acinus in males. The type I acinus was agranular and showed slight morphologic changes during feeding. The presence of cells with ultrastructural features characteristic of epithelia involved in the secretion of hyperosmotic fluids supports the hypothesis that these acini secrete hygroscopic saliva during questing stages to absorb water from an unsaturated atmosphere. There were five granular cell types (a, b, c₁–c₃) in type II acinus, three granular cell types (d, e, and f) in type III acinus, and one granular cell type (g) in type IV acinus. The cells a, d and e secreted most of their granules early in feeding and are considered to be cement precursors. The b and c cells appeared to synthesise and secrete their products throughout feeding and so are likely to secrete anticoagulants, enzymes and other pharmacologically active agents required during feeding. The interstitial cells, which were insignificant in acinar types II, III and IV of unfed ticks, became more distinct during feeding. The type III acinus in females showed remarkable cell transformations, during the course of feeding. The ablumenal interstitial cells of type III acinus, in females formed a basal labyrinth of extraordinary complexity by interdigitating with the basolateral membranes of transformed f cells to form a network of extracellular channels to excrete fluid during feeding. There was an enormous increase in the secretory granules of g cells as the feeding advanced. The secretory granules were released by a process of exocytosis, by direct fusion with the apical membranes and through channels connecting several granules.     

Autophagy-related genes from a tick, Haemaphysalis longicornis

Ticks are gorging-fasting organisms;(1) their life cycle is characterized by alternate off-host (starvation) and on-host (meal) conditions. Their generation time is estimated in several years and many ticks spend more than 95% of their life off the host. They seem to have a unique strategy to endure the off-host state for a long period. Thus, we focused on autophagy, which is induced by starvation and is essential for extension of the lifespan,(2-4) and hypothesized that ticks also have a system of autophagy to overcome the starved condition. Recently, we showed the existence of a homologue of an ATG gene, ATG12, and its expression pattern from nymphal to adult stages in a three-host tick, Haemaphysalis longicornis. The expression level of HlATG12 was downregulated at the beginning of feeding and was highest at 3 months after engorgement. In addition, the HlAtg12 protein was localized to the region around granule-like structures within midgut cells of unfed adults. These results indicate that HlATG12 functions during unfed stages. Here, a potential role of autophagy in unfed ticks is discussed with regard to reports in other animals, such as yeast, mammal, and fruit fly.     

Surface irregularity induced-tunneling behavior of the formosan subterranean termite

Subterranean termites forage from place to place by tunneling through soil. In order to examine termite-tunneling responses to external factors, we designed a square arena that contains five introduction chambers connected with narrow paths, which was filled with sand. Triangle-shaped indentation with width W and height H was provided as surface irregularity on the sand facing the introduction chambers by using templates. After termites were introduced into the chambers, we measured elapsed time, τ, for a tunnel to reach the point at 3 mm away from the apex of the irregularity. We found that for W = 0 mm (the absence of irregularity), termites (Coptotermes formosanus Shiraki) did not tunnel for 7 h, whereas, for W = 1, 2, 3, and 4 mm, they exhibited the tunneling behavior within 20 min. The result indicated that the presence of surface irregularity is essential to induce termite tunneling. In addition, we found that W was correlated with τ, whereas H did not influence τ. This was briefly discussed in the context of individual movement behavior.     

Microhabitat selection by two reptile ticks at their parapatric boundary

The parapatric boundary between the reptile ticks. Aponomma hydrosauri and Amblyomma limbatum, near Mt Mary, South Australia, coincides with a vegetational ecotone. Samples of litter from each side of the tick boundary showed considerable heterogeneity, but there were consistent differences in the composition and quantity of litter. North of the boundary, where Amb. limbatum is found, there is less litter and an absence of Eucalyptus leaves in the litter. Since the litter retards evaporative water loss, and Ap. hydrosauri is less tolerant of dehydration, reduced litter quantity may reduce the fitness of colonizing Ap. hydrosauri. In laboratory and field trials, larvae and nymphs of both tick species chose litter microhabitats in preference to bare soil, with the exception of fed Ap. hydrosauri larvae which, in trials with litter from north of the boundary, rarely moved from the bare soil. This behaviour may also reduce the fitness of colonizing Ap. hydrosauri. The two species adopted different positions in the litter. Ap. hydrosauri were predominantly found on the surface, at the interface between soil and litter as fed larvae, and on or under the surface as nymphs. Amb. limbatum larvae were more often buried under the surface, while nymphs were found mostly in the litter above the soil surface. Hypotheses to explain the maintenance of the parapatric boundary are discussed in the light of these results.     

Field sampling of unfed adults ofAmblyomma hebraeum Koch

No methods have previously been available for the field sampling of unfed adults ofAmblyomma hebraeum. We released 2000 unfed adults into a 5-m² area in mopane woodland in southeastern Zimbabwe. The ticks sought shelter beneath the debris on the soil surface but emerged and became active in host seeking when stimulated. Effective stimuli were the odours of cattle and sheep and high concentrations of carbon dioxide from dry ice. The ticks did not respond to low concentrations of carbon dioxide, vibrations generated by stamping the ground with poles, or two volatile components of ox breath known to attract tsetse flies. The presence of humans in the release area also had little or no effect in stimulating the ticks. As adults ofA. hebraeum are large and conspicuous they can easily be collected from the soil surface, either manually or using a vacuum device, after stimulation by cattle or the release of high concentrations of carbon dioxide.     

Reason for fluctuation in the demographic structure of the taiga tick population (Ixodidae) in dark conifer-leaf forests of Kemchug Upland

It was found out, that mean ratio of unfed taiga ticks Ixodes persulcatus having a life cycle of 3-5 and 6 years counts respectively (N = 6) 33.6, 52.8, 13.2 and 0.4%. Data on absolute number of I. persulcatus individuals in the process of onthogenesis is given. It is shown, that mortality of different unfed stages increases from larva to imago. In the autumn-winter period, the mean ratio of eliminated individuals counted 16% of larvae, 20% of nymphs, and 38 of imago. The mortality in the spring-summer period caused mainly by the deficit of hosts and counted for these stages 3, 82 and 98% respectively. Engorged ticks successfully undertake unfavorable conditions both in the spring-summer and autumn-winter periods. Their mean mortality in the period from feeding to moulting does not exceed 12%.     

Cloning and characterization of an autophagy-related gene, ATG12, from the three-host tick Haemaphysalis longicornis

Ticks are obligate hematophagous ectoparasites with a life cycle characterized by a period of starvation; many ticks spend more than 95% of their life off the host. Autophagy, which is the process of bulk cytoplasmic degradation in eukaryotic cells, is induced by starvation and is essential for extension of the lifespan. Therefore, we hypothesized that autophagy also occurs in ticks; however, there has been no report on autophagy-related (ATG) genes in ticks. Here, we show the homologue of an ATG gene, ATG12, and its expression pattern from the nymphal to adult stages in the three-host tick Haemaphysalis longicornis. The sequence analysis showed that H. longicornis ATG12 (HlATG12) cDNA is 649bp, has a 411bp ORF coding for a 136-amino acid polypeptide with the carboxy-terminal glycine residue, and has a predicted molecular mass of 15.2kDa. Moreover, RT-PCR revealed that HlATG12 was downregulated at the beginning of feeding, upregulated after engorgement, and downregulated again after molting. The expression level of HlATG12 was highest at 3 months after engorgement. By immuno-electron microscopy, it was demonstrated that HlAtg12 was localized to the region around granule-like structures within midgut cells of unfed adults. In conclusion, HlATG12 might function during unfed and molting stages.     

Ant predation on different life stages of two Australian ticks

Laboratory colonies of Rhytidoponera ants were allowed to prey on the fed and unfed stages of the Australian ticks Aponomma hydrosauri and Amblyomma limbatum. The unfed tick stages had a higher survival than the fed stages. The ants took longer to handle the adult ticks than the nymphs and longer to handle the nymphs than the larvae. The ants also took longer to handle the unfed than fed nymphs, but longer to handle the fed than unfed females. As well as the differences between the tick stages, there was a species effect, with the ants taking longer to handle A. limbatum, and with that tick species having a higher survival than A. hydrosauri after ant predation. These stage and species differences may influence the tick population dynamics. This revised version was published online in November 2006 with corrections to the Cover Date.     

极端干旱区沙土掩埋对凋落物分解速率及盐分含量动态的影响

极端干旱区由于降水稀少, 植被盖度低, 太阳辐射强烈, 以及土壤稳定性差, 导致其凋落物周转不同于非干旱区。为探究极端干旱区凋落物分解规律, 该研究利用凋落物分解袋法, 以塔克拉玛干沙漠南缘沙漠-绿洲过渡带优势物种花花柴(Karelinia caspia)、骆驼刺(Alhagi sparsifolia)和胡杨(Populus euphratica)凋落叶为研究对象, 设置不同的沙土掩埋处理: 地表、2 cm和15 cm埋深, 以模拟自然条件下凋落物分解环境, 测定分解过程中凋落物质量和水溶性盐的变化特征。结果表明: 极端干旱区凋落物分解速率与凋落物初始碳(C)含量、氮(N)含量、C:N和木质素含量的关系与非干旱区存在较大差异, 在地表处理下, 木质素含量越高, 质量损失越快。不同分解环境下凋落物质量和水溶性盐损失具有显著差异, 与15 cm埋深相比, 地表和2 cm埋深处理显著增加了凋落物的质量损失和水溶性盐总量损失。地表处理增加了凋落物分解前期的水溶性盐溶解量。该研究表明, 极端干旱区凋落物分解的驱动机制具有独特性, 由于降水稀少, 土壤微生物的活性较低, 掩埋深度不是驱动凋落物分解的主要因素, 极端干旱区凋落物的分解主要受其他非生物过程如太阳光辐射的影响。     

The infection of ixodid ticks collected from humans with the tick-borne encephalitis virus in Tomsk city and its suburbs

The ticks Ixodes persulcatus and I. pavlovskyi collected from people visited gardens and suburban forests have been examined by the IFA methods on the presence of tick-borne encephalitis virus (TEV). It was established, that the most part of ticks collected from humans belongs to I. persulcatus, despite the fact that I. pavlovskyi dominates on the territory of the city and its suburbs. TEV infection was registered more often in fed ticks, in comparison with those without signs of preceding feeding. Infected specimens constituted 9.4% of 53 I. persulcatus individuals examined. In I. pavlovskyi 3.7% of 27 examined specimens were infected. In fed ticks (41 specimens of I. persulcatus and 20 I. pavlovskvi) the rates of infected specimens were higher and constituted 48.8 and 35%, respectively. In suburban forests, as well as in the city, I. persulcatus was found to attack humans more often than I. pavlovskyi. TEV infection was recorded in 12.7% of 220 unfed I. persulcatus specimens examined and in 41.6% of 142 fed individuals. In I. pavlovskyi, 6.6% of unfed and 25% of fed specimens was infected (33 unfed and 24 fed specimens were examined). Thus, in all territories examined, percent of infected specimens was significantly lower in I. pavlovskyi, and in fed ticks of both species, TEV was recorded more often.     

Major surface protein 1a effects tick infection and transmission of Anaplasma marginale.

Anaplasma marginale, an ehrlichial pathogen of cattle and wild ruminants, is transmitted biologically by ticks. A developmental cycle of A. marginale occurs in a tick that begins in gut cells followed by infection of salivary glands, which are the site of transmission to cattle. Geographic isolates of A. marginale vary in their ability to be transmitted by ticks. In these experiments we studied transmission of two recent field isolates of A. marginale, an Oklahoma isolate from Wetumka, OK, and a Florida isolate from Okeechobee, FL, by two populations of Dermacentor variabilis males obtained from the same regions. The Florida and Oklahoma tick populations transmitted the Oklahoma isolate, while both tick populations failed to transmit the Florida isolate. Gut and salivary gland infections of A. marginale, as determined by quantitative PCR and microscopy, were detected in ticks exposed to the Oklahoma isolate, while these tissues were not infected in ticks exposed to the Florida isolate. An adhesion-recovery assay was used to study adhesion of the A. marginale major surface protein (MSP) 1a to gut cells from both tick populations and cultured tick cells. We demonstrated that recombinant Escherichia coli expressing Oklahoma MSP1a adhered to cultured and native D. variabilis gut cells, while recombinant E. coli expressing the Florida MSP1a were not adherent to either tick cell population. The MSP1a of the Florida isolate of A. marginale, therefore, was unable to mediate attachment to tick gut cells, thus inhibiting salivary gland infection and transmission to cattle. This is the first report of MSP1a being responsible for effecting infection and transmission of A. marginale by Dermacentor spp. ticks. The mechanism of tick infection and transmission of A. marginale is important in formulating control strategies and development of improved vaccines for anaplasmosis.     

Susceptibility of <Emphasis Type="Italic">Amblyomma variegatum</Emphasis> ticks to acaricides in Ghana

The susceptibility of unfed and fed stages of larvae, nymphs and adult females of Amblyomma variegatum ticks were tested using Shaws filter paper dip method against four acaricides; chlorfenvinphos and dioxathion, chlorfenvinphos, gamma benzene hexachloride and amitraz at four different concentrations including the recommended dose rates. Based on their lethal concentrations (LC₅₀ & LC₉₀) chlorfenvinphos and dioxathion combined and chlorfenvinphos alone placed first and second, respectively, in all stages except at the unfed nymphal stage where gamma benzene hexachloride topped with a LC₅₀ of 0.001629, while chlorfenvinphos and dioxathion combined and chlorfenvinphos alone had LC₅₀ of 0.001794 and 0.002258, respectively. Amitraz appeared to have a quick knock-down effect on larvae and nymphs but at the recommended dose rate, showed no mortality of the ticks at that stage. However, at a concentration of 0.040%, amitraz showed a 100% inhibition of oviposition and hatching of laid eggs. Gamma benzene hexachloride produced only 66% inhibition of oviposition while chlorfenvinphos and dioxathion combined and chlorfenvinphos alone produced 100% inhibition of oviposition at their recommended dose rates. Fed nymphs were more susceptible than the unfed nymphs. Eggs laid by engorged female ovipositing ticks, applied with gamma benzene hexachloride, hatched.This revised version was published online in May 2005 with a corrected cover date.     

Development, survival, fecundity and behaviour of Haemaphysalis (Kaiseriana) longicornis (Ixodidae) at two locations in southeast Queensland.

The free-living stages of the tick Haemaphysalis (Kaiseriana) longicornis were studied at Mt Tamborine (526 m altitude) and Amberley (25 m altitude) in southeast Queensland between 1971 and 1980. Data are presented on the number of eggs and larvae produced, the moulting success of engorged larvae and nymphs and the survival and behaviour of unfed larvae, nymphs and adult females. Temperature, moisture, daylength, grass length and age of unfed ticks were investigated as sources of variation in development rates, fecundity or survival. At Mt Tamborine the life cycle was well synchronized with the seasons to produce one generation per year. At Amberley higher temperatures accelerated development rates and would have delayed diapause, so disrupting the life cycle.     

Pathogenicity of entomopathogenic fungi to different developmental stages of Rhipicephalus sanguineus (Acari: Ixodidae).

The pathogenicity of 4 species of entomopathogenic fungi (Hyphomycetes species: Beauveria bassiana, Metarhizium anisopliae, Metarhizium flavoviride, and Paecilomyces fumosoroseus) to various developmental stages of Rhipicephalus sanguineus (Acari: Ixodidae) ticks was compared under laboratory conditions. The most virulent isolate, M. anisopliae-108, caused 92-96% mortality to unfed larvae and nymphs on day 7 postinfection (PI) and 100% mortality to unfed adults and engorged females on day 21 PI. The pathogenicity of M. anisopliae-108 to engorged larvae and nymphs was lower--82.6 and 60%, respectively. All tested B. bassiana, M. flavoviride, and P. fumosoroseus isolates were significantly less virulent (P < 0.05) or avirulent toward most life stages of R. sanguineus. The M. anisopliae and M. flavoviride isolates also prevented or reduced the ability of the ticks to lay eggs several days before their deaths. Female ticks infected by the fungi achieved only 11.3-60.8% of their egg-laying capacity compared with the controls.     

华南典型树种凋落叶的野外分解和溶解性有机质溶出动态

选用华南亚热带地区常见阔叶树种木荷和针叶树种湿地松的新近凋落叶,在野外分解0、30、60、90、150、210、240、365 d,分析溶出的溶解性有机质(DOM)浓度、组成和性质的变化,以及对土壤可溶性有机碳(DOC)的影响.结果表明: 随着分解的进行,尽管木荷叶片的DOM浓度高于松针,但是2种凋落叶DOM浓度、性质和物质组成变化规律一致;2种凋落叶的DOM浓度均呈下降趋势,芳香化程度和分子量增大,富里酸、腐殖酸类物质逐渐增多,可降解的简单芳烃蛋白(如酪氨酸)逐渐减少.在分解初期,DOM主要由亲水中性和酸性部分组成,易分解、易迁移,对表层土壤DOC影响不显著;在分解后期,DOM主要为腐殖酸和富里酸类物质,吸附性强,表层土壤DOC浓度显著下降.     

沙地樟子松人工林叶片-枯落物-土壤氮磷化学计量特征

为揭示沙地樟子松人工林N、P分配格局及化学计量特征,以呼伦贝尔沙地、科尔沁沙地、毛乌素沙地不同龄组(中龄林、近熟林和成熟林)沙地樟子松人工林为研究对象,测定分析其叶片、枯落物和土壤N、P含量及化学计量比.结果表明: 研究区3个龄组沙地樟子松人工林叶片、枯落物和土壤N、P含量分别为0.17～49.02和0.11～3.01 g·kg^-1,N/P为0.51～19.74,均表现为叶片＞枯落物＞土壤,且N含量和N/P在3个组分间存在显著差异,叶片P含量显著高于枯落物和土壤.不同地区或林龄对沙地樟子松人工林各组分N、P含量及N/P有一定的影响,但地区和林龄的交互作用对沙地樟子松人工林各组分N/P无显著影响.随着林龄的增加,沙地樟子松各组分N、P含量也增加,在成熟林达到最大值,而N/P没有表现出明显的规律.沙地樟子松人工林N、P含量及N/P在3个组分间呈显著正相关关系.呼伦贝尔沙地和科尔沁沙地樟子松叶片N/P在14.53～15.57,说明这两个地区沙地樟子松人工林的生长可能受N、P的共同限制;毛乌素沙地樟子松叶片N/P在18.56～19.71,说明该地樟子松人工林生长可能受P限制,且林龄对沙地樟子松N、P养分限制的影响不显著.建议在沙地樟子松人工林抚育管理时,依据当地实际情况适当添加N肥或P肥,以提高沙地樟子松林的生产力.研究结果有助于进一步了解N、P在沙地樟子松人工林叶片-枯落物-土壤系统中的相互作用与制约规律,并为沙地樟子松人工林经营管理提供科学依据.     

中亚热带植被恢复阶段植物叶片、凋落物、土壤碳氮磷化学计量特征

为揭示植被恢复过程中生态系统的养分循环机制及植物的生存策略, 根据亚热带森林群落演替过程, 采用空间代替时间方法, 以湘中丘陵区地域相邻、环境条件基本一致的檵木(Loropetalum chinensis) +南烛(Vaccinium bracteatu) +杜鹃(Rhododendron mariesii)灌草丛(LVR)、檵木+杉木(Cunninghamia lanceolata) +白栎(Quercus fabri)灌木林(LCQ)、马尾松(Pinus massoniana) +柯(Lithocarpus glaber) +檵木针阔混交林(PLL)、柯+红淡比(Cleyera japonica) +青冈(Cyclobalanopsis Glauca)常绿阔叶林(LCC)作为一个恢复系列, 设置固定样地, 采集植物叶片、未分解层凋落物和0-30 cm土壤样品, 测定有机碳(C)、全氮(N)、全磷(P)含量及其化学计量比, 运用异速生长关系、养分利用效率和再吸收效率分析植物对环境变化的响应和养分利用策略。结果表明: (1)随着植被恢复, 叶片C:N、C:P、N:P显著下降, 而叶片C、N、P含量和土壤C、N含量、C:P、N:P显著增加, 其中LCC植物叶片C、N含量, 土壤C、N含量及其N:P, PLL植物叶片P含量, 土壤C:P显著高于其他3个恢复阶段, 各恢复阶段植物叶片N:P > 20, 植物生长受P限制; 凋落物C、N、P含量及其化学计量比波动较大。(2)凋落物与叶片、土壤的化学计量特征之间的相关关系较弱, 叶片与土壤的化学计量特征之间具有显著相关关系, 其中叶片C、N、P含量与土壤C、N含量、C:N (除叶片C、N含量外)、C:P、N:P呈显著正相关关系; 叶片C:N与土壤C、N含量、C:P、N:P, 叶片C:P与土壤C含量、C:N、C:P, 叶片N:P与土壤C:N呈显著负相关关系。(3)植被恢复过程中, 叶片N、P之间具有显著异速生长关系, 异速生长指数为1.45, 叶片N、P的利用效率下降, 对N、P的再吸收效率增加, LCC叶片N利用效率最低, PLL叶片P利用效率最低而N、P再吸收效率最高。(4)叶片N含量内稳态弱, 而P含量具有较高的内稳态, 在土壤低P限制下植物能保持P平衡。植被恢复显著影响叶片、凋落物、土壤C、N、P含量及其化学计量比, 叶片与土壤之间C、N、P含量及化学计量比呈显著相关关系, 植物通过降低养分利用效率和提高养分再吸收效率适应土壤养分的变化, 叶片-凋落物-土壤系统的N、P循环随着植被恢复逐渐达到“化学计量平衡”。