Mineral nutrition of cacao (Theobroma cacao L.)

Summary Two experiments on cacao seedlings grown in sand culture are described, the first of which was concerned with variations in the levels of nitrogen, phosphorus, potassium, calcium, magnesium and sulfur (the macronutrient experiment) and the second dealing with variations in the levels of iron, copper, zinc, boron, manganese, and molybdenum (the micronutrient experiment).Many of the deficiency symptoms obtained were similar to those reported in the literature and they have not been described again. However, additional information is provided for symptoms of phosphorus, potassium, calcium, iron, manganese, copper, boron, and molybdenum deficiencies.The effects of all treatments on the dry weights of leaves, stems, and roots are presented. The effects of the macronutrient treatments on the levels of nitrogen, phosphorus, potassium, calcium, and magnesium in the leaves of eight month old plants and the effects of micronutrient treatments on the levels of nitrogen, phosphorus, potassium, calcium, magnesium, iron, manganese, copper, zinc, boron, molybdenum, sodium, and aluminium in the leaves of eleven-month-old plants are presented and discussed.     

Spent medium analysis for liquid culture micropropagation of <Emphasis Type="Italic">Hemerocallis</Emphasis> on Murashige and Skoog medium

Residual nutrients from Murashige and Skoog medium were analyzed following a 5-wk multifactor experiment. Plant density, sugar concentration, and plant growth regulators (benzyladenine and ancymidol) were examined using four genotypes of daylily (Hemerocallis) to determine which factors most influenced nutrient use. Active nutrient uptake was observed for 11 nutrients (potassium, sodium, copper, phosphorus, iron, calcium, magnesium, manganese, boron, sulfur, and zinc) with lower concentrations in spent medium than in the tissue water volume (fresh-dry mass expressed as mL H₂O). Two patterns of nutrient use were visualized by correlative analysis of nutrient uptake. Greatest growth lowered plant nutrient concentrations of potassium, sodium, phosphorus, iron, and copper in all genotypes, and luxuriant uptake was indicated with least growth. Potassium, sodium, iron, and copper concentrations in plant dry matter were equal to or exceeded what is observed in vigorously growing nursery plants. However, phosphorus concentration in plant dry matter was low enough to be considered deficient when compared to Hemerocallis plants in nursery production. With a second group of nutrients (calcium, magnesium, manganese, and boron), the genotype, “Barbara Mitchell” lacked active uptake and was deficient. Calcium concentration was low in all plants compared to Hemerocallis grown under nursery conditions (“Barbara Mitchell” was the lowest concentration) despite active uptake by the other three genotypes—“Brocaded Gown,” “Mary’s Gold,” and “Heart of a Missionary.” Magnesium concentration in these three genotypes was low enough in vessels with greatest growth to question its adequacy at high densities. Increased sucrose in medium reduced the dry matter concentrations of all tested nutrients. Plant growth regulators had less impact on nutrient use than genotype and plant density. Nutrient uptake may be an important physiological component of genotypic variation.     

Nutrient Cycling in an Age Sequence of Western Washington Douglas-fir Stands

The cycling of nitrogen, phosphorus, calcium, magnesium andpotassium in a series of western Washington Douglas-fir [Pseudotsugamenziesii (Mirb.) Franco] stands ranging in age from 9 to 95years has been described. The stands were of relatively lowproductivity being limited by low nitrogen. The content of nitrogen,phosphorus, magnesium and potassium in tree foliage all tendedto stabilize at about 40 years whereas calcium continued toincrease. The content of all nutrients in the wood continuedto increase with stand age. Nitrogen in the forest floor accumulatedconstantly at about 5.7 kg ha^–1 year^–1 and thistogether with the above-ground tree accumulation meant about10.5 kg ha^–1 year^–1 nitrogen was immobilized. Calciumalso increased with time in the forest floor with age whereasthe other nutrients were fairly constant after about 30 years.Understorey nutrient content reached a peak at about 20 years,while understorey litter-fall was significant throughout theage sequence. Internal redistribution, especially of nitrogen,represented an increasingly greater proportion of stand requirementwith increasing stand maturity. Pseudotsuga menziesti (Mirb.) Franco, Douglas-fir, biomass, litter-fall, nutrient content, nutrient cycling     

NUTRIENT AND WATER RELATIONS OF THE MISTLETOE PHORADENDRON LEUCARPUM (VISCACEAE): HOW TIGHTLY ARE THEY INTEGRATED?

Tissue mineral concentrations of calcium, copper, iron, magnesium, manganese, nitrogen, phosphorus, potassium, sodium, and zinc, and long-term water-use efficiency (as estimated by δ¹³C±) were measured in the mistletoe Phoradendron leucarpum and its hosts to investigate the relationship between nutrient concentrations and water economies of the parasite and hosts. The mistletoe had mineral concentrations 0.97 to 2.88 times greater than the hosts. Mean long-term mistletoe water-use efficiency (-27.89±) was comparable to mean long-term host water-use efficiency (-27.69±) and generally greater than the water-use efficiency reported for most mistletoe species. Differences between mistletoe and host water-use efficiency ranged from +0.65 to -0.75± and were more similar to mistletoes found growing on nitrogen-fixing host species as reported in other studies. Mistletoe nutrient concentrations, in particular nitrogen, were not related to changes in mistletoe water-use efficiencies. Nutrient: calcium ratios indicated that mistletoes acquire nutrients in excess of that which can be delivered by the host xylem. These data are discussed relative to the passive vs. active theories of nutrient uptake.     

Radial and vertical variation of wood nutrients in Bornean tropical forest trees

Nitrogen, phosphorus, potassium, calcium, and magnesium concentrations in woody tissue are poorly documented, but are necessary for understanding whole-tree nutrient use and storage. Here, we report how wood macronutrient concentrations vary radially and along the length of a tree for 10 tropical tree species in Sabah, Malaysia. Bark nutrient concentrations were consistently high: 2.9–13.7 times greater than heartwood depending on the nutrient. In contrast, within the wood both the radial (sapwood vs. heartwood) and vertical (trunk bottom vs. trunk middle) variation was modest. Higher concentrations in sapwood relative to heartwood provide empirical support for wood nutrient resorption during sapwood senescence. Dipterocarp species showed resorption rates of 25.3 ± 7.1% (nitrogen), 62.7 ± 11.9% (phosphorus), and 56.2 ± 12.5% (potassium), respectively, while non-dipterocarp species showed no evidence of nutrient resorption in wood. This suggests that while dipterocarps have lower wood nutrient concentrations, this family is able to compensate for this by using wood nutrient resorption as an efficient nutrient conservation mechanism. In contrast to other nutrients, calcium and magnesium tended to accumulate in heartwood. Wood density (WD) showed little vertical variation along the trunk. Across the species (WD range of 0.33 to 0.94 mg/cm³), WD was negatively correlated with wood P and K concentration and positively correlated with wood Ca concentration. As our study showed exceptionally high nutrient concentrations in the bark, debarking and leaving the bark of the harvested trees on site during logging operations could substantially contribute to maintaining nutrients within forest ecosystems.     

Biogeography and variability of eleven mineral elements in plant leaves across gradients of climate, soil and plant functional type in China

Understanding variation of plant nutrients is largely limited to nitrogen and to a lesser extent phosphorus. Here we analyse patterns of variation in 11 elements (nitrogen/phosphorus/potassium/calcium/magnesium/sulphur/silicon/iron/sodium/manganese/aluminium) in leaves of 1900 plant species across China. The concentrations of these elements show significant latitudinal and longitudinal trends, driven by significant influences of climate, soil and plant functional type. Precipitation explains more variation than temperature for all elements except phosphorus and aluminium, and the 11 elements differentiate in relation to climate, soil and functional type. Variability (assessed as the coefficient of variation) and environmental sensitivity (slope of responses to environmental gradients) are lowest for elements that are required in the highest concentrations, most abundant and most often limiting in nature (the Stability of Limiting Elements Hypothesis). Our findings can help initiate a more holistic approach to ecological plant nutrition and lay the groundwork for the eventual development of multiple element biogeochemical models.     

The effect of iron supply on the yield and composition of leaves of tomato plants

Summary This paper describes a factorial experiment which was designed to elucidate the effect of total iron supply and rate of iron supply on the yield of tomato plants. Information was also obtained on the effect of iron supply and leaf age on the concentration of a number of nutrient elements in the leaf tissue.Increasing the total iron supply increases the yield, and the iron concentration in the leaves, while the manganese, phosphorus, sodium, calcium and magnesium decrease in concentration. The young leaves have a lower concentration of iron, manganese, potassium, sodium, calcium and magnesium than the older leaves, while the reverse is true of nitrogen and phosphorus. A slow rate of iron supply decreases the yield and sodium concentration in the leaves but increases the manganese and phosphorus concentrations.Interaction between total iron supply and leaf position affects the manganese, potassium, calcium and magnesium concentrations in the leaves, while the interaction between total-iron level and rate of iron supply affects the phosphorus and calcium concentration in the leaves.     

Effects of Aluminium on Honeylocust (Gleditsia triacanthos L.) Seedlings in Solution Culture

Honeylocust (Gleditsia triacanthos L.) seedlings were grownin solution culture at pH 4.0) with 50, 150, 600 and 1500 mmolm^–3 aluminium. All levels of aiuminium reduced the sizeand weight of roots, shoots and leaves with the exception ofroot elongation at 50 mmol m^–3 Al. Aluminium content ofroots was 50 to 100 times that of shoots. With increasing concentrationof aluminium, aluminium content of leaves and roots increasedexponentially while a linear increase was observed for stems.The nutrient content of seedlings was improved in 50 mmol m^–3where increases in shoot calcium, magnesium, and phosphorusconcentrations were observed. Aluminium concentrations greaterthan 50 mmol m^–3 reduced shoot nutrient content. Presenceof aluminium increased the root phosphorus and calcium levelsbut had no effect on potassium and magnesium concentrations.Results show that honeylocust is an aluminium sensitive treespecies whose growth may be reduced by high soil Al levels. Key words: Aluminium toxicity, Gleditsia triacanthos, nutrient solution     

Nutrient requirement for UASB process: a review

In literature, numerous studies regarding nutrients dosing are available in UASB reactors. Various nutrients and tracemetals used in UASB studies include nitrogen, phosphorous, yeast extract, magnesium, potassium, calcium, sulphur, iron, aluminium, zinc, nickel, cobalt, molybdenum, manganese, copper, boron, selenium, citrate, resazurine, EDTA, and hydrochloric acid. Often, there is a wide variation in the composition and quantification of nutrients and tracemetals, used to supplement the influent feed in UASB reactors. In this paper, nutrients and tracemetals formulations used in different studies on UASB reactors are compiled and critically analysed. In order to have uniformity, concentrations of nutrients and tracemetals have been converted into equivalent terms as g nutrients or tracemetals/g of waste COD. The ranges for this ratio for all the constituents of nutrients and tracemetals are given. Besides, the ranges associated with minimum start-up periods are also obtained. It is hoped that the information presented in this study would facilitate the nutrients dosing in UASB reactors.     

Rainfall pattern and nutrient content influences on African elephants’ debarking behaviour in Samburu and Buffalo Springs National Reserves,Kenya

The magnitude of debarking by elephants was investigated in Samburu and Buffalo Springs National Reserves. About 1617 plants were monitored for debarking intensities for 6 months spanning through dry and wet seasons. Debarking indices ranged from no debarking at all during the wet months to complete stem girding at the height of the dry season. A negative correlation was found between rainfall and debarking indices. It was hypothesized that nutrient content of the bark influenced the magnitude to which trees were debarked. Bark samples were collected from least, moderate and intensely debarked plants throughout the 6 months. These were analysed for calcium (Ca), sodium (Na), phosphorus (P), magnesium (Mg), potassium (K), nitrogen (N), iron (Fe), copper (Cu), manganese (Mn) and zinc (Zn). Significant positive correlations were found between debarking intensity and each of the nutrients N [crude protein (CP)], P, K and Zn. Bark was found to be richest in CP and Calcium. Neutral detergent fibre content was on average 67%. Monthly variations in nutrient composition were minimal. Acacia elatior, the most preferred species had significantly higher quantities of each of the four elements N, P, K and Zn than Acacia tortilis, the second most preferred woody species.     

Nutrient concentrations in two savannah grass species and their implications for grazers

The diversity of grazers on savannahs exerts pressure on the grass to provide nutrition. In this study the concentrations of major and trace animal nutrients in above ground grass tissue as influenced by grazing were examined. Two palatable grass species grazed by both livestock and wild grazers were sampled: Brachiaria nigropedata and Eragrostis lehmanniana. Samples of grass tissue were collected from widely spread sampling sites in both livestock and wildlife grazing sites, at the end of the growing season. Although B. nigropedata had higher nutrient concentrations, there was general covariance in nutrient levels in the two grasses. The covariance indicated that the respective sites generally had location context-dependent low or high nutrient concentrations in all grasses. The adequacy of the nutrient concentrations for grazer nutrition was assessed using literature-sourced minimum and maximum tolerable limit requirements of cattle and the high population gazelle-like species as illustration. The results showed that, in these nitrogen-limited savannahs, the grass generally met grazer requirements in terms of crude protein, potassium, molybdenum, nickel and zinc but that it had deficiencies in phosphorous, sodium, and boron, as well as grazing intensity-related localised toxic levels of potassium, phosphorous, magnesium, iron and manganese. The deficiencies and toxicities were inferred to result in abnormalities in grazer physical condition and reproduction. Given the importance of livestock in semiarid savannahs, these effects on livestock grazers were inferred as threatening human food security. The study shows that high grazing intensity can raise the concentrations of some animal nutrients in above ground grass tissue to levels above grazer tolerable limits. Grazing rotation to reduce grazing intensity is recommended as a proactive grazing management strategy to limit the effects.     

Polyphenolic and biological activities of leaves extracts of Argemone subfusiformis (Papaveraceae) and Urtica urens (Urticaceae)

Nowadays there is a resurgence of interest in wild plants for their possible medicinal value in diets, since some epidemiological studies have demonstrated their effectiveness against important diseases. Generally, foods of plant origin contain many bioactive compounds, proteins, energy, vitamins and specific minerals; in addition, the popular wild plant species provide fibre, essential fatty acids and enhance the taste and colour in diets. We studied the nutritional and medicinal potential of leaves of A. subfusiformis and U. urens, collected in Alice, South Africa in November 2006. To asses this we analyzed the phytochemical, antioxidant and antibacterial activities of leaves in acetone, methanol and water extracts, using standard analytical methods. The proximate analysis showed that the leaves of both plant species contained appreciable percentages of moisture; ashes; carbohydrates; crude proteins, lipids and fibres. Elemental analysis of macro and microelements showed higher values for U. urens that contained in decreasing order: iron > manganese > zinc > copper > calcium > potassium > nitrogen > magnesium > phosphorus > sodium, for A. subfusiformis resulted in iron > zinc > manganese > copper > calcium > potassium > nitrogen > magnesium > phosphorus > sodium. Besides, the chemical composition showed higher concentration of alkaloids, saponins and phytates in A. subfusiformis. The extracts also caused 1, 1-diphenyl-2-picrylhydrazyl (DPPH) and 2, 2-azinobis-3 ethylbenzothiazoline-6-sulfonic acid (ABTS) radical scavenging activities which were comparable to those of ascorbic acid. The extracts of both plants contained appreciable levels of polyphenols and also caused varied inhibition of some bacterial strains used in this study. When the nutrient and chemical constituents of these plants were compared with recommended dietary allowance (RDA) values, the results revealed that the leaves contain an appreciable amount of nutrients, minerals, and phytochemicals and low levels of toxicants. Since the plants also exhibited some level of antibacterial activities, their use for medicinal purposes is to some extent being justified.     

Mineral and phytochemical influences on foliage selection by the proboscis monkey (Nasalis larvatus)

Primate diets may be influenced by the phytochemical composition of potential food resources. This study examined the protein, fiber, and mineral content of leaves from plant species used as food resources by the proboscis monkey (Nasalis larvatus) in Tanjung Puting National Park, Kalimantan Tengah, Indonesia. In addition, leaf samples were collected from nonfood resources based upon relative density in previously established vegetation plots. Higher concentrations of crude protein, phosphorus, potassium, and zinc were found in young leaves. Mature leaves had higher levels of magnesium, manganese and calcium compared to young leaves. Leaves from species used as food resources were significantly higher in protein (9.9% vs. 8.5%) and lower in fiber (30.5% vs. 40.0% acid detergent fiber) than the leaves from species not used as food sources. Food resources also contained higher concentrations of phosphorus and potassium; nonfood resources contained higher concentrations of calcium and manganese. Proboscis monkeys appear to follow the typical colobine trend: a diet relatively high in protein and low in digestion inhibitors (fiber). Little previous work has examined mineral content as a selection factor in primate food choices. Leaves from species used as food resources contained less calcium (0.35%), iron (34 mg/kg) and phosphorus (0.15%) than dietary standards established for captive nonhuman primates, but values were similar to those established for ruminant herbivores. Other mineral concentrations (potassium, magnesium, manganese, sodium, and zinc) met or exceeded recommended levels for both primates and ruminants. Comparison of diverse nutrients in native diets may provide insight into suitability of domestic animals models for providing adequate diets in captive feeding situations, as well as help define resource limitations in natural environments. Am. J. Primatol. 41:117–128, 1997. © 1997 Wiley-Liss, Inc.     

Current and Future Transgenic Whole-Cell Biosensors for Plant Macro- and Micronutrients

From the soil, plants take up macronutrients (calcium, magnesium, nitrogen, phosphorus, potassium, sulfur) and micronutrients (boron, chloride, cobalt, copper, iron, manganese, molybdenum, nickel, selenium, and zinc). In acidic soils, aluminum can interfere with nutrient uptake. There is a need for improved diagnostic tests for these soil-derived minerals that are inexpensive and sensitive, provide spatial and temporal information in plants and soil, and report bioavailable nutrient pools. A transgenic whole-cell biosensor detects a stimulus inside or outside a cell and causes a change in expression of a visible reporter such as green fluorescent protein, and thus can convert an invisible plant nutrient into a visible signal. Common transgenic whole-cell biosensors consist of promoter-reporter fusions, auxotrophs for target analytes that are transformed with constitutively expressed reporters, riboswitches and reporters based on Forster Resonance Energy Transfer (FRET). Here, we review transgenic plant biosensors that have been used to detect macronutrients and micronutrients. As plant-based biosensors are limited by the requirement to introduce and optimize a transgene in every genotype of interest, we also review microbial biosensor cells that have been used to measure plant or soil nutrients by co-inoculation with their respective extracts. Additionally, we review published transgenic whole-cell biosensors from other disciplines that have the potential to measure plant nutrients, with the goal of stimulating the development of these diagnostic technologies. We discuss current limitations and future improvements needed, and the long-term potential of transgenic whole-cell biosensors to inform plant physiology, improve soil nutrient management, and assist in breeding crops with improved nutrient use efficiency.     

北京地区耕地碳酸盐褐土不同农作物氮及灰分元素生物循环的比较研究

京郊耕地碳酸盐褐土小麦、大麦、玉米、高梁、花生五种农作物氮及灰分元素生物循环的研究结果表明：1．不同作物存留、归还、吸收不同元素数量的差别很大。豆科作物花生以对钙、镁吸收量大为其显著特征;禾本科作物吸收较高数量硅、铁、铝、锰、钠。在四种禾本科作物中,高梁吸收较高数量氮、磷,钾、钙、镁、硅,大麦吸收较高数量钠、硫和锰。2．不同作物以根、茬形式对11种元素和灰分向土壤中的归还有三种情况：1)低度归还类型：氮磷钾,归还比例一般低于10%;2)中度归还类型：镁钙硫硅钠和灰分,归还比例介于10—30%;3)高度归还类型：铁铝和锰,归还比例一般为30%。 3．不同农作物生物循环特点对研究农田生态系统元素平衡、耕地土壤肥力变化和形成过程以及施肥、耕作制度改革,都有重要意义。     

Plant genotype differences in the uptake,translocation, accumulation,and use of mineral elements required for plant growth

Plant genotypes differ in their uptake, translocation, accumulation, and use of mineral elements. Examples of genotype differences to iron, nitrogen, phosphorus, potassium, calcium, magnesium, manganese, boron, copper, zinc, and molybdenum are discussed. Current knowledge is sufficient to indicate that many crop plants can be improved for the efficient use of mineral elements and better adaptation to mineral stress conditions.     

The Levels of Manganese, Sodium, Aluminium, Iron, Boron, Zinc, and Copper in the Leaves of Cacao (Theobroma cacao L.): As Affected by Shade, Fertilizer, Irrigation, and Season

The results are given of chemical analyses of leaves collectedover a period of one year from cacao trees in a field trialreceiving fertilizer, shade, and irrigation treatments. Thegreatest variation in concentration of the elements under studywas found to be due to the time of year at which the leaveswere collected. The lowest concentrations of zinc and copperwere found in the leaves collected during the peak of the mainseason crop. The lowest concentrations of manganese, aluminium,and iron were found in leaves collected about a month afterthe end of the main harvest. The levels of most elements rosesharply after the end of the main harvest. The levels of mostelements rose sharply after reaching a minimum, and the maximumlevels were usually reached before completion of flowering forthe next main season crop. The effects of the treatments onthe nutrient levels in the leaves were generally small. Thefertilizer treatment tended to increase the levels of manganese,aluminium, iron, sodium, and zinc and to decrease the levelof boron. The shade treatment tended to increase the levelsof sodium, copper, and zinc and to decrease the levels of manganese,aluminium, and iron. The irrigation treatment had no substantialeffect on the level of any element. The levels of iron, manganese,and boron, were found to be positively correlated with the levelof aluminium.     

Resorptions of 10 mineral elements in leaves of desert shrubs and their contrasting responses to aridity

Aims We aim to investigate variations in the resorption efficiencies of 10 mineral nutrients [i.e. nitrogen (N), phosphorus (P), potassium (K), magnesium (Mg), calcium (Ca), manganese (Mn), zinc (Zn), aluminum (Al), iron (Fe) and copper (Cu)] in leaves of desert shrubs and to explore effects of aridity on resorption efficiency of these nutrients.     

Inefficient nitrogen resorption in genets of the actinorhizal nitrogen fixing shrubComptonia peregrina: physiological ineptitude or evolutionary tradeoff?

Nutrient resorption was measured in an actinorhizal nitrogen-fixing shrub,Comptonia peregrina, for five years in the understory of a deciduous oak forest in Rhode Island, USA. Mean resorption of nitrogen was extremely inefficient (11%) compared to most deciduous species (50%+), yet resorption of phosphorus was efficient (53%) and comparable to other species. Of the seven additional nutrients studied, only copper (6%) and zinc (10%) were resorbed from senescing leaves. Resorption of nitrogen (5%–20%) and phosphorus (40%–71%) varied significantly among years. Copper was resorbed from leaves in three years and accreted into leaves in two years. Five-year resorption means differed among individual genets by as much as a factor of 2.5 for nitrogen, and 1.3 for phosphorus. Resorption of nitrogen, copper, and zinc were highly correlated, yet resorption of phosphorus remained autonomous from other nutrients. The ecophysiological tradeoffs inComptonia which have resulted in the cooccurence of actinorhizal nitrogen fixation, inefficient nitrogen resorption, and efficient phosphorus resorption suggest that plant nutrient status does have an impact on resorption efficiency and that the evolution of nutrient conservation strategies is nutrient-specific.     

温带荒漠灌丛和藓类结皮对土壤养分的贡献

灌丛与生物土壤结皮镶嵌分布是温带荒漠常见的地表景观之一,二者的发育均显著影响了地表土壤养分的空间分布特征及循环过程。然而,灌丛和生物土壤结皮对荒漠土壤表层养分的影响存在怎样的差异,二者对养分变化的贡献度如何尚不清楚。因此,选择中国北方典型温带荒漠古尔班通古特沙漠为研究区,以荒漠中的优势灌丛膜果麻黄(Ephedra przewalskii)灌丛和生物土壤结皮发育高级阶段的藓类结皮的结皮层及结皮下层土壤为研究对象,采集不同微生境(裸露地、灌丛下)的裸沙与藓类结皮的土壤样品。为探究不同微生境下不同土层碳、氮、磷和钾养分变化特征,测定了不同土层土壤有机碳(SOC)、全氮(TN)、全磷(TP)、全钾(TK)、速效氮(AN)、速效磷(AP)和速效钾(AK)含量。结果表明:(1)相较于裸露地,灌丛显著提高了藓类结皮不同土层有机碳、全氮和全钾的含量和裸沙全磷的含量,降低了灌丛下藓类结皮土壤全磷的含量。(2)对于速效养分而言,与裸沙相比,裸露地藓类结皮覆盖降低了土壤速效氮含量,增加了速效磷和速效钾含量;而灌丛下藓类结皮覆盖提高了土壤速效氮和速效钾的含量,但降低了速效磷的含量。 (3) 相关性分析显示,在0-2 cm土壤中速效磷与速效氮呈现显著负相关,而在2-6 cm速效磷与速效氮呈现显著正相关(P<0.01)。(4)贡献度分析表明,土壤中灌丛效应对养分的贡献(42.54%)要远大于藓类结皮的贡献(2.43%),但二者交互作用却降低了除速效氮以外的其他土壤养分含量。综上,灌丛、藓类结皮覆盖和土层深度变化对土壤碳、氮、磷和钾养分均具有显著影响(P<0.05),但三者间的交互效应对养分的影响不显著(P>0.05)。相对于裸露地,灌丛与藓类结皮的覆盖均对表层土壤碳、氮、磷和钾养分具有促进作用,且随着土层深度的下降,土壤养分含量呈现显著的下降趋势(P<0.05)。因此,在荒漠生态系统中耐旱灌丛与生物土壤结皮这两个最重要的斑块单元联合调控了微尺度土壤养分的空间异质性变化。