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Soil Ecology

by: Mitchell Schulist

Soil Ecology NRE 430

Mitchell Schulist
GPA 3.51


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This 19 page Class Notes was uploaded by Mitchell Schulist on Thursday October 29, 2015. The Class Notes belongs to NRE 430 at University of Michigan taught by Staff in Fall. Since its upload, it has received 30 views. For similar materials see /class/231581/nre-430-university-of-michigan in Natural Sciences at University of Michigan.

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Date Created: 10/29/15
Introduction An ecosystem is a three dimensional space occupied by interacting abiotic and biotic components through a period of time The abiotic components are made up of climate physiography and soil where as the biotic components consist of plants animals and microbes These elements are interconnected thus giving rise to ecosystem structure and function Barnes et al 1998 Soil is an abiotic component of landscape ecosystems The soil resource contains physical chemical and biological properties These properties interact and control soil processes and ultimately ecosystem productivity nutrient cycling and energy ow Zak 2002 Ecologically soils provide substrate for plant development and microbial activity habitat for living organisms the supply and ltration of water and the cycling of energy and nutrients throughout the ecosystem Brady amp Weil 1999 The formation of soil depends on regional climate the activities and processes of organisms the composition of the parent material topography and the length of time in which physical chemical and biological ecosystem processes have occurred Zak 2002 These interconnected factors shape soil properties and processes which include soil texture rates of leaching and weathering pH nutrient status and moisture regimes The ecosystems located in northern and southern Lower Michigan have contrasting regional climate and physiography Albert et al 1989 The varying climate generates differential temperatures and precipitation regimes which affect ecosystem properties such as evapotranspiration growing season length rates of decomposition and degree of leaching and weathering The physiography of Michigan has been highly in uenced by glaciers that retreated 10000 to 16000 years ago Albert et al 1989 Over time the composition of physiography in uenced by climate weathering and biota has given unique characteristics to the soils of northern and southern regions of Lower Michigan Albert et al 1989 The makeup of the parent material along with climate and time have strongly in uenced soil formation and as a result have de ned the soil texture available water content soil pH and nutrient status that exist today The objectives of this study were to compare and contrast ecosystem productivity of two ecosystems located in southeastern and northwestern Lower Michigan Soils are the foundation on which ecosystems form and therefore soil properties and processes provide insight into ecosystem function and structure Zak 2002 When ecosystem components including soil are analyzed and treated as separate entities these two ecosystems are seemingly different due to dissimilar climate and parent material However when this information is combined and viewed as smaller dependent parts of their respective larger systems the interworkings of each ecosystem can be understood and productivity can be compared Barnes et al 1998 Description of Study Sites The Mixed Oak ecosystem MO is located in southern Lower Michigan in Washtenaw County In this region of Michigan the average annual temperature is 93 C and average precipitation MaySeptember is 380 mm The growing season lasts for 163 days and elevation is 840 ft Albert et al 1989 This ecosystem is found on a moraine derived from calcareous fine textured glacial till In presettlement times this ecosystem was dominated by American beech F agus grandz39folia and Sugar maple Acer saccharum However after clearing for timber Northern red oak Quercus rubra assumed the dominant position in the overstory The Northern Hardwoods ecosystem NH is located in the Manistee National Forest in Wexford County Michigan At this location average annual temperature is 67 C and average precipitation MaySeptember is 400 mm Elevation is 1070 ft and the length of the growing season is 115 days Albert et al 1989 The landform is a sandy moraine formed from glacial till parent material This forest was clearcut in the early 1900 s and is currently dominated by Sugar maple Acer saccharum Methods Field Data Collection Data was collected on September 25 and October 5 2002 at the Mixed Oak ecosystem and the Northern Hardwoods ecosystem respectively A 15 x 30 m plot 045 ha was constructed at each ecosystem and soil pits were established Percent slope aspect and position in the landscape and drainage class were estimated Soil horizons were delineated and depth structure texture color percent coarse fragments pH and boundary classi cations were taken for each Twelve cores 10 cm depth 1 inch diameter and two bulk density cores 10 cm depth 2 inch diameter of mineral soil were randomly collected from within the plot Two 0 horizon samples were collected using 025m2 quadrats All samples were stored for further laboratory analysis Woody and herbaceous vegetation was characterized in the overstory understory and groundcover Trees greater than 10 cm in diameter at breast height DBH were counted and their diameters measured Basal area per hectare and relative dominance was calculated Soil Physical Properties Soil texture was determined using the Hydrometer method Because the velocity of a particle settling out of solution is directly related to the size of its squared radius larger particles like sand settle out rst and smaller particles like silt settle out after Clay pa1ticles due to their platelike structure remain suspended Hydrometer readings were taken twice 40 seconds and 2 hours after plunging From these readings the percentages of sand silt and clay were calculated Soil texture was estimated with the Textural Triangle Available water content AWC is the amount of water that plants are theoretically able to uptake from the soil Speci cally it is the amount of water held on soil particles between the permanent wilting point 15 MPa and eld capacity 001 MPa To nd AWC the pressure plate method was employed From this AWC was found by calculating the difference between water held at field capacity and permanent wilting point Soil Chemical Properties Soil pH measures the concentration of H ions in soil solution Soil was combined with deionized water and CaClz Two glass electrodes calibrated to pH 4 and pH 7 respectively measured the pH of the upper 10 cm of soil at each ecosystem The Wet Combustion method was used to calculate organic matter content of the soil Sulfuric acid and l M potassium dichromate were used to oxidize the organic carbon in soil samples and a spectrophotometer was used to estimate the amount of reduced dichromate ion A spectrometer was used to analyze percent transmittance of light through the soil solution and 5 carbon standards Using this information linear regression was performed for the carbon content of standards versus log transmittance of standards Organic carbon content of the samples was calculated using the equation of the regression line and the known log transmittances From this percent organic carbon and percent organic matter were calculated assuming organic matter is 50 carbon The amount of exchangeable hydrogen and aluminum ions in each ecosystem s soil was analyzed by ooding the exchange complex with other ions allowing measurement for H and Al ions in solution KCl was added to airdry soil the mixture was filtered and Phenolphthalein indicator was added This solution was titrated with NaOH until the concentration of H equaled the concentration of OH39 A blank solution of KCl was also titrated to analyze the amount of acidity it contained Total acidity from H and Al combined was the difference in the amount of NaOH used in the titration of the soil mixture and the KCl blank To analyze the amount of exchangeable base cations Ca2 Mg K and Na an Inductively Coupled PlasmaOptical Emission Spectrometer ICPOES was used NH4Cl was added to airdry soil and the mixture was ltered This solution was then analyzed by the ICP OES to determine concentration of base cations in the soil The cation exchange capacity of each soil was calculated as the sum of all base cations and total acidity Base saturation was calculated as the percentage of base cations on the CEC Soil Biological Properties Laboratory analyses were used to estimate soil microbial biomass Water was added to soil samples to bring them to eld capacity One sample was used as a control and the other was fumigated with chloroform for 20 hours After this period the chloroform was evacuated from the fumigated samples and were inoculated with soil from the control samples All samples were placed into airtight mason jars and incubated at room temperature 22 C for 13 days After incubation gas samples were extracted from both the fumigated and control jars and analyzed using a gas chromatograph to determine the amount of carbon dioxide produced by soil microbes Microbial biomass was calculated by subtracting the carbon dioxide of the control sample from the fumigated sample and diViding by a correction factor Microbial respiration rate per gram of soil was calculated by using the amount of carbon dioxide from the control sample From this specific microbial respiration rate C02 respiredmicrobial biomass was found To calculate nitrogen mineralization KCl was added to the incubated control soil samples used in the soil biomass analysis which extracted cations from the CEC Via mass action The mixtures were filtered and analyzed using a Rapid Flow Analyzer for ammonium and nitrate amounts Net N mineralization was calculated by diViding the difference of incubated ammonium and nitrate and control ammonium and nitrate by the number of days incubated Similarly Net N nitrification was calculated by diViding the difference of incubated nitrate and control nitrate by number of days incubated The ratio of carbon respired to nitrogen mineralized a measure of organic matter quality was calculated by dividing the microbial respiration rate described above and the net nitrogen mineralization rate Biomass and nitrogen pools were calculated for the aboveground forest oor and belowground portions of the ecosystem The aboveground pool includes all biomass and nitrogen accumulated in the boles branches and leaves of standing trees Equations based on speci c tree species and tree diameters were used to calculate the biomass Nitrogen content was calculated using known nitrogen concentrations in the aboveground pools of speci c tree species Forest oor biomass was calculated by weighing forest oor samples Nitrogen content was calculated using known nitrogen concentrations of litter found in similar ecosystems Belowground biomass of organic matter was estimated for the top 10 cm of the soil by using the bulk density and organic matter content of our soil calculated previously Nitrogen content of the soil was calculated using a known nitrogen concentration of similar ecosystems for the first 10 cm of soil Results Field Data Results The drainage classes for M0 and NH are moderately well drained and well drained respectively MO was located on the midslope of a moraine with a westsouthwest aspect with a slope of 5 NH is positioned on the upperslope of a morainal ridge with an eastern aspect and slope of 3 MO and NH soil profile information is summarized in Appendices l and 2 respectively MO has a thinner A horizon than NH Both ecosystems have E horizons however it is considerably deeper at MO NH has 38 cm of B horizon Bh B51 and B52 where as MO has a B1 horizon with a width of 20 cm The C horizon of NH is found deeper in the soil pro le 90 cm than MO 72 cm The overstory tree species found at MO in order of greatest to least basal area per hectare include Quercus rubra 2472 mzha Acer rubrum 792 mzha Acer saccharum 096 mzha Sassafras albidum 091 mzha Carya ovata 032 mzha Ostrya virginiana 027 mzha and Amalanchier arborea 02l mzha Total basal area per hectare at this ecosystem is 3530 mzha Graph 1 displays the relative dominance of each overstory tree species with their respective values Q rubra is the most dominant tree species followed by A rubrum The MO understory and ground cover includes species such as Prunus sp Vitis rz39parz39a A rubrum O virginiana Fraxz39nus americana Rubus sp and Viburnum acerz39folium Mixed Oak Relative Dominance 8000 7000 m 0 o 0 5000 4000 3000 2000 Relative Dominance 1000 090 077 060 Quercus Acerrubrum Acer Sassafras Carya ovata Ostrya Amalanchier rubra saccharum abidum viginiana arborea Tree Species Graph 1 Graphical representation including values of the relative dominance of the MO ecosystem The overstory tree species at NH in order of greatest to least basal area per hectare include Acer saccharum 2077 mzha Tilia Americana 594 mzha Quercus rubra 507 mzha and Ostrya virginiana 046 mzha Total basal area per hectare at this site is 3224 mzha A saccharum accounts for 64 of the total dominance followed by T Americana with 18 relative dominance Graph 2 NH understory and ground cover species are A saccharum O viriginana T Americana Aaliantum sp Allium sp Arialia nualicaclis Carex sp Osmorhiza chilensis Prunus serotina anal Trillium sp brthern Hardwoods Relative Dorrinenoe Waive Danier 55313139 p CL v Treeaaecies Graph 2 Graphical representation including values of the relative dominance of the NH ecosystem Physical Soil Properties Based on laboratory results MO has a sandy loam texture borderline loam and NH has a loamy sand texture borderline sand Graph 3 compares the soil textural components for M0 and NH As evidence by the graph NH has a higher sand component than MO where as MO has a greater percentage of clay than NH AWC for M0 and NH is 45 045 cm3 watercm3 soil and 34 034 cm3 watercm3 soil respectively Soil TexlueBradown UNIXedm EmmiW Particles sand s39lt clay Particle Size Gas Graph 3 Graphical comparison of the 3 soil particle size classes for the MO and NH ecosystems Chemical Soil Properties Table 1 provides soil pH values and percent organic matter for the two ecosystems MO has higher pH values in water and calcium chloride than NH and is therefore is more basic in the upper 10 cm of soil Organic matter was similar for M0 and NH with 351 and 359 respectively Soil pH Organic Matter pH in H20 pH in Old Mixed Oak 616 509 351 Northern Hardwoods 564 458 359 Table 1 Soil pH and organic matter for the MO andNH ecosystems Values based on class average data Acidity and Cation Exchange Capacity CEC information is presented in Table 2 Total acidity in both ecosystems was similar and relatively low The CEC and base saturation was greater in the MO ecosystem than in the NH ecosystem Total Acidity CEC Base Saturation cmollkg cmolkg Mixed Oak 016 924 1018 Northern Hardwoods 008 547 985 Table 2 Total acidity CEC and Base saturation for the MO andNH ecosystems Biological Soil Properties Data regarding the microbial communities and their activities at each ecosystem are reported in Tables 3 and 4 The microbial biomass at the MO and NH were similar however MO had a higher microbial respiration rate and speci c respiration rate Net nitrogen mineralization and carbon respired to nitrogen mineralized are slightly greater in NH however net nitri cation is the same in both ecosystems Microbial Biomass Microbial Respiration Rate Specific Respiration Rate g Cm mggday mggday Mixed Oak 2720 3781 19419 Northern Hardwoods 2626 2801 122 86 Table 3 Microbial biomass respiration rate and specific respiration rate for the MO andNH ecosystems Net N Mineralization Net N Nitri cation Carbon Respired to g NmZday g NmZday N itrogen quot Ratioquot Mixed Oak 053 002 648 Northern Hardwoods 061 002 668 Table 4 Net nitrogen mineralization and nitrification and CN ratio for the MO and NH ecosystems Ecosystem biomass pools are summarized in Table 5 and graphically compared in Graph 4 1n MO and NH the aboveground biomass component contained the most carbon followed by the belowground component and the forest oor with the least carbon Overall MO had more Values based on class average data biomass accumulation in the aboveground component however NH had more carbon accumulation in the belowground and forest oor components Total biomass was greater in MO Aboveground Biomass Forest Floor Biomass Belowground Biomass Total Biomass Mg C m Mg C m Mg C m Mg C m Mixed Oak 30158 450 3366 33974 Northern Hardwoods 22780 1019 4775 28574 Table 5 Aboveground forest floor and belowground biomass for the MO and NH ecosystems DWaZlCK EMITI EmI Bdnax s Biorrms N9 Cl39a W Forest Floor mgum Graph 4 Graphical comparison of the aboveground forest floor and belowground biomass pools for the MO and NH ecosystems Table 6 and Graph 5 display ecosystem nitrogen pools for M0 and NH In MO and NH most nitrogen resides in the 39 39 g A r of the J t The aboveground component contains the second greatest amount followed by the forest oor with the least nitrogen The NH ecosystem has more nitrogen than MO in the belowground component and the forest oor and MO has a greater nitrogen content located in its aboveground component than NH Overall NH has the greater total nitrogen Aboveground Nitrogen Forest Floor Nitrogen Belowground Nitrogen Total N kg N m kg N m kg N m Mixed Oak 75491 6080 258930 340501 Northern Hardwoods 61960 12528 305900 380388 Table 6 Aboveground forest floor and belowground nitrogen for the MO and NH ecosystems Ecosystem Ntrogen Pools DWaZlCK EMITI EmI Bdnax s Ntrogan Cortert kg Mn Abwegum Forest Floor mgum Graph 5 Graphical comparison of the aboveground forest floor and belowground nitrogen pools for the MO and NH ecosystems Discussion Physical Properties NH has an overall deeper soil profile thicker A and B horizons due to the texture climate and length of the growing season In this sandy soil water percolates downward at a faster rate than MO which has a clayey Bt horizon that slows water percolation Thus leaching is faster at NH The growing season is shorter and colder in the NH ecosystem Therefore with less evapotranspiration and fewer days in which water is used by plants there is greater leaching and greater depth to the soil The soil at MO is a typical alfisol with high base saturation and a B1 horizon Conversely NH is a spodosol that is acidic coarser in texture and has accumulated humus iron and aluminum in its B horizons Zak 2002 The soil texture of the two ecosystems is a function of the parent material on which it formed MO has more clay and silt because the parent material is ne textured till with high clay and silt content NH has a greater proportion of sand and much less silt and clay because it formed from sandy till The compositional differences in parent material at each ecosystem are due to the material mixed in with the glacial ice Because MO and NH are at different locations different materials were present in the glacier when the parent material was laid 10000 to 16000 years ago Zak 2002 The higher AWC in MO can be explained by soil texture MO had higher percentages of clay and silt and much less sand than NH This texture allows for greater surface area enabling greater water adsorption to soil particles Zak 2002 The soil at the NH ecosystem had a relatively low AWC because compositionally it was mostly sand However the soil does contain relatively high available water for plant uptake due to the cooler climate more precipitation during the growing season and high soil organic matter in the rooting zone Soil organic matter greatly increases the amount of water that the soil can hold and is especially important in sandy soils Brady amp Weil 1999 Chemical Properties The lab and field analysis of soil pH conclusively show that MO soil is more basic than NH in all horizons This is a result of parent material and time in uencing soil texture and thus base saturation The parent material of MO is a calcareous till which contains greater concentrations of base cations than the sandy parent material of NH The loamy texture is a 13 result of the weathering of the calcareous parent material through time The high clay and silt content allow for greater surface area on which the abundant base cations adhere Zak 2002 The high base saturation is a measure of the base cation capacity of the CEC which is greater than NH The MO CEC contains more base cations than hydrogen and aluminum ions and more nutrients are available to plants when compared to NH as indicated by the higher total acidity at NH The contents of the CEC re ect the contents of the soil solution and thus there are fewer of these acidic ions in solution at MO Zak 2002 Soil organic matter SOM was similar for both ecosystems and therefore the rate of decomposition is similar Field observations support this conclusion the O horizons at each site were similar thickness and both lacked an Oa horizon High microbial biomass and low forest oor biomass at each site are also indications of rapid decomposition This fast turnover rate of litter breakdown supplies plants with necessary nutrients However this result is seemingly unintuitive for NH Generally in colder areas such as Northern Michigan breakdown of litter is slow because the lower temperatures negatively affect microbial activity Brady amp Weil 1999 However closer inspection of the quality of the litter at each site provides insight to rates of decay Bohlen et al 2001 Carreiro et al 1999 Maithani et al 1998 Quercus leaves contain more lignin than Acer leaves and therefore decompose more slowly Finzi et al 1998 Geng et al 1993 Q rubra and A saccharum are the dominant tree species at MO and NH respectively However MO contains Acer spp and NH contains Q rubra at intermediate dominance levels MO decay rates are most likely only slightly slowed and NH rates only slightly increased by leaf lignin concentration The carbon respired to nitrogen ratio CN verifies the lower quality litter at MO The CN ratio is slightly lower for M0 than NH signifying that the MO litter is harder for microbes to break down Nitrogen mineralization is controlling the nitrogen balance rather than nitrogen immobilization at M0 to a higher degree than NH Because the leaves have more difficult carbon compounds to break down lignin more microbes cleave nitrogen from organic molecules to obtain energy than assimilating nitrogen into their biomass Zak 2002 Because the CN ratio is higher at NH the litter is higher quality and is more easily broken down Nitrogen is incorporated into microbial biomass more so than MO and less available for plant uptake Zak 2002 Seneviratne et al 1999 Both ecosystems exhibit high CBC and base saturation However MO has more base cations and less hydrogen and aluminum ions on the CEC thus has a higher base saturation than NH There was experimental error in the calculation of base saturation due to negative acidity The base saturation for both ecosystems should be lower than reported but MO should theoretically be higher than NH MO has a high base saturation due to high organic matter and clay content and the calcareous till on which the soil was derived NH has a relatively high base saturation due to high soil organic matter and ultimately the rapid decomposition of its litter layer The high base holding capacity of organic matter is extremely important for acidic clay poor soils Johnson 2002 Brady amp Weil 1999 Johnson et al 1997 Pregitzer et al 1983 Biological Properties Quality as in uenced by leaf litter biochemistry and quantity of SOM soil texture temperature and matric potential affect microbial biomass Bohlen et al 2001 Zak 2002 Both ecosystems exhibit similar microbial biomass For M0 the high SOM and loamy texture thus high AWC and warmer temperatures allow for high microbial biomass High microbial biomass at NH is attributed to the high quantity and quality of the SOM which retains soil moisture However MO has a higher microbial respiration rate and specific respiration This indicates that MO microbes are not as efficient at converting carbon into biomass but rather respire more carbon dioxide This is another indication of the poorer litter quality at MO relative to NH 15 Microbes spend more energy breaking down litter and little is left over for incorporation into microbial biomass at MO Also at NH the types of microbes have probably evolved to be more efficient at incorporating carbon into biomass due to poorer site conditions such as low pH and cold climate Fungi are good examples of efficient microbes under these conditions Zak 2002 Nitrogen mineralization is greater at NH than at MO although the figures are very similar The sites have similar decomposition rates SOM and microbial biomass Greater lignin content at MO suggests that it should have greater N mineralization due to carbon limitations However other explanations suggest the contrary Higher pH at MO results in a more diverse microbial community in comparison to NH thus more types of carbon can be broken down Brady amp Weil 1999 Nitrogen nitrification is similar and minimal in both ecosystems Specific bacteria nitrify nitrogen compounds and have specific habitat requirements Brady amp Weil 1999 NH has too low a pH for these organisms to ourish The nitrification rate was expected to be higher in MO than what the data indicate due to higher pH and AWC and warmer temperature than the NH ecosystem Zak 2002 Finzi et al 1998 Ecosystem productivity is a function of soil texture and water availability Host et al 1988 Overall there is more biomass at MO and thus it is more productive than NH MO had more nutrients and water available for plant uptake due to the loam texture in uencing the high base saturation and AWC However NH has a great deal of biomass as well and is relatively highly productive due to its high SOM easily decomposable leaf litter and efficient microorganisms A great amount of nutrients are available here as well as indicated by the high base saturation and a high belowground nitrogen pool higher than MO As for water content the cooler climate allows for less evaporation and transpiration in comparison to M0 NH has higher belowground biomass that is most likely due to texture Because the soil is so sandy the plants must allocate more carbon to root growth to get required water and nutrients which are more limiting in this ecosystem in comparison with the clayey basic soil of MO NH also has more biomass on the forest oor than MO One might expect to see the opposite because Quercus leaves are relatively difficult to decompose However NH may exhibit slightly slower decomposition than MO due to the lower soil pH and cooler temperatures However a more likely explanation is the Summer 2002 drought The drought tolerance of Acer saccharum dominant tree species at NH is lower than Quercus rubra dominant tree species at MO Auge amp Moore 2002 Auge et al 1998 Tschaplinski 1998 Loewenstein amp Pallardy 1997 and thus more leaves fell earlier at NH than at M0 M0 has more aboveground biomass than NH Overall there are better site conditions at the MO ecosystem due to the higher AWC base saturation higher pH greater clay content and longer growing season at this site Comparatively there is more nitrogen in the forest oor and belowground pools of NH than MO most likely due to the greater biomass accumulated in these pools in the NH ecosystem Bohlen 2001 suggest that lower tree growth as measured by basal area and higher litter accumulation at higher elevations due to lower AWC causes lower plant and microbe competition for nitrogen thus resulting in a larger pool in the forest oor The large belowground nitrogen pool matches that of Finzi et al 1998 that found the mineral soil beneath A saccharum dominant at NH had greater nitrogen concentration than the mineral soil beneath Q rubra dominant at MO There is more nitrogen in the aboveground pool of MO than NH because MO has more biomass in this pool Overall NH contains more total nitrogen than MO but has lower total biomass Nitrogen is high at this site because Acer leaves contain more nitrogen than do Quercus leaves Geng et al 1993 and thus through leaf litter vegetation on this site significantly in uences soil nitrogen quot 39 quotquotJ 39 quot 39the 39 39 g 39 pool contains the most nitrogen but there is relatively low biomass in the belowground component of the ecosystem This demonstrates the importance of belowground activities on nutrient cycling Conclusion Not one soil component can solely explain the high productivity at these ecosystems Ecosystem properties and processes are linked to one another through various pathways It is the interconnectedness of these properties that must be understood in order to explain the high productivity of both ecosystems In the case for M0 and NH climate and physiography are largely responsible for this These soilforming factors control soil moisture and available nutrients At MO soil moisture was high due to the loamy soil texture despite the warmer temperature and westsouthwest aspect At NH the gentle eastern slope with high growing season precipitation and low evapotranspiration due to lower temperatures and aspect microclimate effects and high organic matter kept the soil moisture high making it available for plant uptake The high nutrient availability at MO was driven by rapid decomposition due to high microbial biomass which in turn depended on the high water availability The loamy texture and high organic matter aided in high base saturation At NH decomposition was also rapid There was high soil moisture despite the sandy texture due to less evapotranspiration and high organic matter The SOM held water that aided in the diffusion of carbon molecules for uptake by microbes Zak 2002 Microbial biomass was high and the leaves were relatively easier to breakdown and contained more nitrogen than the leaf litter at MO All in all high moisture nutrients and microbial biomass due to climate and physiography give these ecosystems their highly productive characteristics Literature Cited Albert DA SR Denton amp BV Barnes 1989 Regional Landscape Ecosystems of Michigan School of Natural Resources and Environment University of Michigan Auge RM amp JL Moore 2002 Stomatal response to nonhydraulic roottoshoot communication of partial soil drying in relation to foliar dehydration tolerance Environmental and Experiemental Botany 47 217229 Auge RM X Duan JL Croker WT Witte amp CD Green 1998 Foliar dehydration tolerance of twelve deciduous tree species Journal of Experimental Botany 49 753759 Barnes BV DR Zak SR Denton amp SH Spurr 1998 Forest Ecology 4g Edition John Wiley amp Sons Inc New York Chapter 2 Bohlen PJ PM Groffman CT Driscoll TJ Fahey amp TG Siccama 2001 PlantSoil Microbial Interactions in a Northern Hardwood Forest Ecology 824 965978 Brady NC amp RR Weil 1999 The Nature and Properties of Soils PrenticeHall Inc New Jersey Pages 434 amp 468 Carreiro MM K Howe DF Parkhust amp RV Pouyat 1999 Variation in quality and decomposability of red oak leaf litter along an urbanrural gradient Biology and Fertility of Soils 30 258268 Finzi AC NV Breemen amp CD Canham 1998 Canopy TreeSoil Interactions within Temperate Forests Species Effect on Soil Carbon and Nitrogen Ecological Applications 82 440446 Geng X J Pastor amp B Dewey 1993 Decay and nitrogen dynamics of litter from disjunct congeneric tree species in oldgrowth stands in northeastern China and Wisconsin Canadian Journal ofBotany 71 693699 Host GE K S Pregitzer CW Ramm DP Lusch and DT Cleland 1988 Variation in overstory biomass among glacial landforms and ecological land units in northwestern Lower Michigan Canadian Journal ofForestResearch 18 659668 Johnson CE 2002 Cation exchange properties of acid forest soils of the northeastern USA European Journal ofSoil Science 53 271282 Johnson CE RB Romanowicz amp TG Siccama 1997 Conservation of exchangeable cations after clearcutting of a northern hardwood forest Canadian Journal of Forest Research 27 859868


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