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'11!" - 11 . 111 11111" llll Ill l lll‘llllllllllllllllllll . 1, _ 3 1293 00103 6700 THESIS LIBRARY Michigan State University This is to certify that the thesis entitled NUTRITIONAL RESPONSE OF WILDLIFE FORAGES T0 MUNICIPAL SLUDGE APPLICATION presented by Henry Campa, III has been accepted towards fulfillment of the requirements for M.S. Fisheries and Wildlife degree in Major professor Date August 18, 1982 0-7639 MS U is an Affirmative Action/Equal Opportunity Institution IVIESI_J RETURNING MATERIALS: Place in book drop to Lian/nuns remove this checkout from n your record. FINES will be charged if book is returned after the date stamped below. ‘I AH? '3 it” ““540- k“ 5 :3 ., Ll NUTRITIONAL RESPONSE OF WILDLIFE FORAGES TO MUNICIPAL SLUDGE APPLICATION By Henry Campa, III A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Fisheries and Wildlife 1982 ABSTRACT NUTRITIONAL RESPONSE OF WILDLIFE FORAGES TO MUNICIPAL SLUDGE APPLICATION By Henry Campa, III Municipal sewage sludge, obtained from the Cadillac, Michigan sewage treatment facility, was applied to study areas in a 4-year-old jack pine (Pinus banksiana) clear-cut. Six 2 ha plots were delineated in the clear-cut, and 3 were randomly selected and treated with approximately 11 metric dry tons/ha of sludge during May, 1980. Samples of selected plant species were collected in summer and late fall. Crude protein, ether extract, ash, phosphorus, Goering and VanSoest fiber analyses, hemicellulose, cellulose, and in yitrg dry matter digestibility values were used to indicate the nutri- tional quality of selected plant species. Samples were also analyzed for concentrations of 13 elements. Changes in annual productivity were monitored on both sludge-treated and control areas. Results indicated total annual productivity was signi- ficantly greater on sludge-treated areas than control areas. Crude protein content for all species (except jack pine) was significantly greater on sludge-treated areas than Henry Campa, III control areas for both seasons. Phosphorus analysis showed only herbaceous species and all fall woody species had significantly greater levels on sludge-treated areas than control areas. Goering and VanSoest fiber analyses, hemi- cellulose, and cellulose analysis indicated a trend of lower fiber levels and thus greater digestibility for plants on sludge-treated areas. Elemental analysis indicated herba- ceous species on sludge-treated areas accumulated significantly greater concentrations of more elements than other plant groups. Levels of these elements are not believed to present any toxicity problems to wildlife. Plant species which showed the greatest increase in nutritional qualities and elements on sludge-treated areas also displayed an increase in production. This was commonly due to: l) depth of the root system of a species and/or 2) speed at which a species can normally assimilate nutrients. These results indicate that sludge application to appropriate forested lands could serve as a viable solution to both sludge disposal and habitat improvement. Future research should focus on the length of time nutritional benefits can be expected with a single application of sludge. ACKNOWLEDGEMENTS This project was cooperatively funded by the U.S. Forest Service, North Central Forest Experiment Station and the Michigan Agriculture Experiment Station. I would like to thank my advisor, Dr. Jonathan Haufler for his guidance, support, and friendship throughout the course of this project. I also want to acknowledge Dr. Harold Prince, Dr. Duane Ullrey, and Dr. Dean Urie for their support and suggestions in preparing my thesis. Assistance in the laboratory was provided by Dave Woodyard, Pat Harkins, Linda Doolittle, Pat Valerance, and Larry Hicks. Advise with data analysis was provided by Dr. Phu Nguyen and Phil Mello. I would like to thank all these people for their help. Acknowledgements are also extended to Dean Urie, John Cooley, Bruce Birr, Julie Patterson, Bill Dunn, and Ray Harris of the U.S. Forest Service, North Central Forest Experiment Station for their advise and help throughout various stages of this project. I would also like to thank Anne Thomas for her help in editing portions of this thesis. A special thanks is extended to Dave Woodyard for his assistance in the field, laboratory and last but most ii importantly his friendship which helped make this project a more memorable experience. I would also like to thank Sue Hazard for typing my thesis. iii TABLE OF CONTENTS Page LIST OF TABLES ....................................... Vi LIST OF FIGURES ...................................... x INTRODUCTION ......................................... l OBJECTIVES ........................................... 7 STUDY AREA DESCRIPTION ............................... 8 METHODS AND MATERIALS ................................ 12 Experimental Design ............................... 12 Sludge Application ................................ 12 Selection of Plant Species to be Sampled .......... 17 Sample Collection ................................. 18 Summer ........................................ 13 Fall .......................................... 19 Sample Preparation ................................ 19 Summer ........................................ 19 Fall .......................................... 20 Chemical Analyses ................................. 20 Statistical Analysis .............................. 20 RESULTS .............................................. 24 Annual Productivity ............................... 24 Crude Protein ..................................... 24 Phosphorus ........................................ 28 Ether Extract ..................................... 28 Ash ............................................... 31 lg vitro Dry Matter Digestibility ................. 31 Cell WaIl Constituents ............................ 31 Cell Soluble Material ............................. 35 Acid-Detergent Fiber .............................. 35 Hemicellulose ..................................... 35 Acid—Detergent Lignin ............................. 37 Cellulose ......................................... 37 iv Elemental Analysis ................................. 37 Season—Treatment Interactions ...................... 55 In vitro Dry Matter Digestibility - Fiber Constituents and Cell Soluble Material ............. 55 DISCUSSION ............................................ 58 SUMMARY AND RECOMMENDATIONS ........................... 78 LITERATURE CITED ...................................... 80 APPENDIX .............................................. 87 LIST OF TABLES Number Page 1 Mean application rates of elements to soils ... 15 2 Mean levels of elements in municipal sludge obtained from Cadillac, Michigan, 1980 ........ 16 3 Above-ground net annual production (kg/ha) <<2 m height on study areas in 1980 ........... 25 4 Regression equations and covariance analysis for net above-ground annual production (g dry wt) per tree height (cm) of jack pine (Pinus banksiana) and cherry (Prunus spp.) in I980 ....................................... 26 5 Comparisons of crude protein content of vegetation from sludge—treated and control areas for samples collected during summer and fall, 1980 .................................... 27 6 Comparisons of phosphorus content of vege- tation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 29 7 Comparisons of ether extract content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 ................... ' ............. 3O 8 Comparisons of aSh content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .................................... 32 9 Comparisons of in vitro dry matter digesti- bility of vegetafion from sludge-treated and control areas for sampled collected during summer and fall, 1980 ................. 33 vi Number 10 ll 12 l3 14 16 17 Comparisons of cell-wall constituents (CWC) and cell-soluble material (CSM) of vegetation from sludge—treated and control areas for samples collected during summer and fall, 1980 ................................ 34 Comparisons of acid-detergent fiber content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 ................................ 36 Comparisons of hemicellulose content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 ................................ 38 Comparisons of acid-detergent lignin content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 ................................ 39 Comparisons of cellulose content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 40 Comparisons of zinc content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 ........ 42 Comparisons of lead content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 43 Comparisons of chromium content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 44 Comparisons of cadmium content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 45 vii Number Page 19 Comparisons of magnesium content of vege- tation from sludge-treated and control areas for samples collected during summer and fall, 1980 ................................ 46 20 Comparisons of potassium content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 47 21 Comparisons of manganese content of vege- tation from sludge-treated and control areas for samples collected during summer and fall, 1980 ................................ 48 22 Comparisons of aluminum content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 49 23 Comparisons of copper content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 50 24 Comparisons of sodium content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 51 1980 .......................................... 25 Comparisons of nickel content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 52 1980 .......................................... 26 Comparisons of iron content of vegetation from sludge—treated and control areas for samples collected during summer and fall, 1980 .................................... 53 27 Comparisons of calcium content of vegetation from sludge-treated and control areas for samples collected during summer and fall, 1980 .......................................... 54 28 F-values of significant interactions between treatments and seasons of element and nutrient contents of vegetation samples ................ 56 viii Number 29 30 Sample correlation coefficients between i3 vitro dry matter digestibility and fiber constituents and cell-soluble material ........ List of vascular plants on the study site ..... ix 87 LIST OF FIGURES Number Page 1 Location of study site near Cadillac, Michigan ..................................... 9 2 Mean precipitation and temperature data by month during study period, 1980 .............. ll 3 Location of sludge-treated and control areas on the study site ............................ l4 INTRODUCTION As the human population has grown, the problem of waste accumulation has also increased. Disposing of large volumes of municipal waste has caused public concern and an increasing demand for environmentally safe methods of waste disposal. Presently, municipal wastewater treatment plants are producing approximately 18 to 91 mil- lion metric tons of sludge (the solids accumulated and concentrated during wastewater treatment) annually in the U.S. (Lang 1981 unpubl. data). With such a large volume of waste, problems are arising in obtaining sites and local public approval for implementing any of the acceptable sludge management alternatives. Currently, the alternatives for disposing of this vast amount of sludge are limited to incineration, landfilling, ocean disposal,enuiland application. If disposal sites are available and liquid sludges are accepted, land- filling may be an economical alternative. Hauling distances, however, may be a limiting factor in imple- menting landfilling as a sludge disposal method. If sites are not available, incineration is the method employed most often if air quality criteria allow it. Ocean disposal is expected to be phased out, since dumpers do not meet the ocean dumping requirements. Due to the limitations of these disposal methods, with rising fuel costs, the land application of sludge is being imple- mented and investigated as a primary method of sludge disposal for the future (Bastian 1977). Because of the growing public desire and need to recycle resources, municipal sewage sludge produced by modern sewage treatment techniques has been applied to agricultural lands, and the effectscniplant quality and productivity studied. By using this means of sludge disposal, society can capitalize on a viable, economical and readily available source of nutrients for crop pro- duction (Urie 1979). Because sewage sludge provides nitrogen and other nutrients, researchers have observed that when sludge is applied to agricultural crops, their productivity and nutritive quality have been enhanced (Blessin and Garcia 1979). In addition to nitrogen, these waste products are a source of phosphorus, calcium, zinc, iron, and sulphur for plants. Because of some of the additional elements present, problems do exist with the land application of sewage sludge. The major concerns with applying sludge to land are: l) aesthetics, 2) the possibility of introducing deleterious chemicals (such as heavy metals) and persistent organic compounds, and 3) pathogenic microorganisms occuring in the human food chain. When sludge is applied to agricultural lands, there is concern that the marketed portion of crops grown on sludge- treated soils will be in direct or indirect contact with the sludge. Despite the fact that both aerobic and anaerobic sewage treatment processes are quite effective in removing pathogens and toxic chemicals, no technique is 100% effective. The possibility of heavy metal ac- cumulation in foodstuffs remains a significant cause for concern (Bitton eE_§l. 1980). Sagik 35 El' (1979) stated that while arsenic, copper, lead, mercury, nickel, selenium and zinc are of concern, cadmium poses the greatest human health risk. They also stated, however, that if control is exerted on domestic sewage at point sources, requiring them to use adequate secondary treatment followed by disinfection, the resultant waste may contain lower levels of potential human pathogens and chemical contaminants. Because of such problems, researchers have sought safer alternative sludge disposal sites that will still benefit from the fertilizer effect. By using forested areas in- stead of croplands, the chance of toxic chemical and pathogen transfer to the human food chain is reduced (Sopper 1973). In addition,tflmaunsightly appearance of sludged vegetation is secluded. A primary goal of applying sludge to forest lands is to use the nutrients available in sludge to manipulate and improve wildlife habitat quality. Several researchers have documented that the application of sludge to forest lands has resulted in a significant increase in tree growth (Roth g5 El. 1979, Urie 1979). Plant responses produced by sludge application are quite similar to those observed with inorganic fertilizers. These comparable results are produced by the nutrients and trace elements contained in sludges and inorganic fertilizers (Braids 3; El. 1980). Urie (1979) stated that superior plant growth on sludge- treated areas can continue to be anticipated because of the resulting nitrogen reservoir in the soil. Researchers who have studied the effects of sewage effluent or sludge application on agricultural crops and forests have found, in all instances, that the nutritive value of vegetation was improved. The nutrient component which consistently increased with waste application was crude protein (Dressler and Wood 1976, Blessin and Garcia 1979). This is a common index of forage quality among researchers (Bailey 1968, Kelsey 35 El. 1973, Groetz 1975, Bayoumi and Smith 1976, Bear 1978). Blessin and Garcia (1979) studied strip mines and found an increase of 2.5% in protein content after sludge was applied. Dressler and Wood (1976) noted crude protein, potassium and phosphorus were significantly higher in plants grown on effluent irrigated sites, whereas crude fiber and calcium were sig- nificantly lower. Thus, studies have shown that with the land application of sewage waste, plant quality and pro- ductivity increase. As plants mature, however, a definite change in their chemical composition occurs, eventually causing nu- trient quality to decline. Sludge application may accelerate this change due to the influx of nutrients causing plants to reach maturity earlier. An increase in fiber components as plants mature is an example of how nutritive quality declines with maturity (Cook and Harris 1950). Van Soest (1965) stated that as the fiber components of plants increased with maturity, forage digestibility declined. The level of digestibility was closely re— lated to the chemical composition of plants. Presently, recommended rates for the land application of sludge are based on the fertilizer value (nitrogen, phosphorus, and potassium) and on the concentrations of trace metals present in sludge. While appropriate levels of nitrogen, phosphorus, and potassium may enhance plant productivity and quality, excessive amounts of metals may produce a negative effect. Sommers (1977a) stated that when zinc, copper, lead, nickel, and cadmium are applied to soils in excessive amounts, plant yield or quality of food or fiber may be impaired. Because municipal waste land application may produce benefits such as improved plant nutritional value, it could be an established waste treatment and habitat im- provement technique for wildlife management (Urie 1979). Other researchers have suggested that organic waste appli- cation may be the beginning of reclamation programs (Lejcher and Kunkle 1973). It may also be part of a productive process like inorganic fertilization. Most importantly, however, land application processes return nutrients to their natural cycles and dispose of waste in a manner in which any long—term adverse impact could be monitored and corrected if necessary (Freshman 1977). Because of such benefits, and the urgent need for a safer method of sludge disposal, the practiceof sludge appli— cation to appropriate forested lands should be researched for consideration as a viable method of habitat improvement. OBJECTIVES The main objective of this study was to determine the effects of a single sludge application on the nutritional quality, in terms of chemical constituents and digesti- bility of selected wildlife forage species and parts of these plant species during summer and fall. A second objective of the study was to determine how sludge appli— cation would affect annual productivity. Production was investigated since the influx of nutrients may significantly increase growth as well as the quality of vegetation. STUDY AREA DESCRIPTION The study area was located in the 8% N% of Section 15, T22N, R10W in Wexford County, approximately 6.4 km north of Cadillac, Michigan (Figure 1). The site was a 20 ha, 5-year old jack pine clear-cut which had also been roller chopped. This site is within the Cadillac Hilly Upland physio- graphic region of Michigan's lower peninsula. The area is in the watershed of the Muskegon River which drains west- ward to the eastern shore of Lake Michigan (Sommers 1977b). The surface formations of the area are primarily a medium altitude outwash plain of stratified sand and gravel deposits. These materials were deposited in the Pleistocene epoch (Sommers 1977b). Soils on the study site were of the Graycalm and Montcalm series. Both soil types are sandy and range from being excessively drained (Graycalm) to well drained (Montcalm). Graycalm soils are very strongly to slightly acidic while Montcalm soils are considered medium to slightly acidic (Corder 1979). The litter layer is poorly developed with no A-2 horizon present, indicated that cultivation may have occurred within the last century. The slope is z Wexford County * Study Site 0 Cadillac Figure 1. Location of study site near Cadillac, Michigan. 10 gradual and to the west, with a low spot in the southwest corner of the site. The climate of Cadillac alternates between continental and semi-marine in character, as the weather patterns change. Lake Michigan influences Cadillac's climate primarily during the late fall and winter months when westerly winds bring increased cloudiness and milder minimum temperatures. Average annual temperature and precipitation for this area are 5.800 and 76 cm respectively. Mean (1928-1969) and 1980 monthly temperature and precipitation data for the study period are given in Figure 2 (Eichmeier 1963, Strommen 1974, Witehell 1980). The existing vegetation consisted of regenerating jack pine, interspersed by clumps of pin cherry (Prunus pennsyl- vanica), black cherry (Prunus serotina), and choke cherry (Prunus virginiana). The ground cover ranged from areas of dense grasses and brambles (Rubus, spp.) to gaps of no vegetation. ll 3O 28 26 24 E; 22 V 20 8 18 -a E 16 .a 14 :0"... .H 12 : "0' o : "b a) - '0 H 10 o o m § 8 °""""m°o_"'o., is/O/K ull'—- o 6 O '%:\°’$ 0.". .01"... g 0'0 4 1""EF 2 20 18 .98""""""0, Illmu Study period a 16 04 , — Mean (1928-1969) o g? " l4 4“:;P 3 12 a? 3 g 10 o g 8 H 6 4 2 M J J A s O N Months Figure 2. Iean percipitation and temperature data by month during study period, 1980. METHODS AND MATERIALS Experimental Design The experimental design was six-2 ha plots located on the study site. Three of these were randomly selected for sludge-treatment in May 1980, 3 years after clear-cutting (Figure 3). Sludge Application During May 1980, non-industrial municipal sewage sludge was applied to treatment areas at a loading rate of ap- proximately 11 metric dry tons/ha. This loading rate was equivalent to 481.5 kg/ha of nitrogen, 571.4 kg/ha of phosphorus, and 4.2 kg/ha of potassium. The U.S. Forest Service has determined that this nitrogen loading rate does not cause ground water levels of nitrate to exceed safe limits. Loading rates of other elements are given in Table 1. All sludge was obtained from the sewage treatment facility at Cadillac, Michigan. It was produced by a bio- logical secondary sewage treatment process. The sludge received approximately 90 days of anaerobic digestion prior to application. The elemental analysis of the sludge ap- plied is shown in Table 2. 12 13 Figure 3. Location of sludge-treated and control areas on the study site. l4 hardWOOds .3225... 05.... \ @ no; HIIIIIIIIIIIIIIIIIh‘ — Eocw - e no; 50.5 32:00 ...-5 no; nouaozlomUEm I .- I mucous—ta... IIIIIIIIIIIIIIIIIIIII n a9< IIIIIIIIIIIIIIIIIIIII Figure 3. 15 Table 1. Mean application rates of elements to soils. Element Application Rate (Kg/ha) Zn 17.0 Cd 0.6 Mn 7.4 A1 59.0 Mg 75.0 Cu 7.3 K 4.2 Ni 0.8 Cr 2.6 Na 17.0 N 481.5 P 571.4 Fe 752.4 Ca 403.7 16 Table 2. Mean levels of elements in municipal sludge obtained from Cadillac, Michigan, 1980. Element ppm Zn 1535 Cd 56 Mn 671 A1 5388 Mg 6788 Cu 665 K 378 Ni 69 Cr 239 Na 1545 N 43800 P 52000 re 6.8 (Z) Ca 3.7 (X) 17 A portable pipeline-rain cannon system was used to distribute the sludge over the ground and litter surface of treatment areas. Sewage sludge was trucked to the study area and stored in a pit, from which it was pumped through irrigation piping, and sprayed on the areas. Each treatment area was divided into a 6 x 6 grid. The rain cannon was placed at the points of intersection within each grid so that sludge was applied relatively evenly to all treatment areas. Sludge application took 11 days. Selection of Plant Species to be Sampled The emphasis of this project was to determine how the land application of sludge affects the nutrient quality of vegetation in general. The plant species to be sampled were selected on the basis of their relative abundance. Vegetation samples were collected during both summer and fall to allow for seasonal comparisons. Summer samples consisted of cherry, jack pine, brambles, orange-hawkweed (Hieracium aurantiacum), sedge (Carex spp.), and panic grass (Panicum virgatum). Fall samples were collected after leaf fall so chemical analyses would yield mean nutrient and element values representative of plant species during the fall and winter. These samples were confined to cherry, jack pine, and brambles, since by late fall herba- ceous species are dead and often under snow cover, therefore are unavailable for wildlife consumption. Other plant 18 species on the study site are listed in the Appendix, Table 30. Sample Collection Summer The summer sampling period was conducted from mid- July to mid-August 1980. A completely randomized sampling design was used to choose the locations of all plots and starting points of transects for vegetation sampling. To determine annual productivity for cherries and jack pine all current annual growth <2 m was clipped from individual trees. Eight trees from each species were randomly selected on each area with 2 individuals selected from each of 4 height strata. The height strata for cherry and jack pine were: 0-30cm, 30—60cm, 60-90cm, >90cm, and 0-60cm, 60-90cm, 90-120cm, and >120cm respectively. To collect bramble and panic grass samples, random plots were located and the vegetation clipped at ground level. Clipped vegetation from 5 plots was combined to compose a single sample for nutrient and element analysis for these species. To collect orange-hawkweed and sedge, 3 transects were established on each area, running from the northern to the southern border. Each transect had 5 sampling points located 28m apart in order to collect vegetation from the entire length of an area. Approximately 100g of vegetation 19 were collected from the closest individual(s) to each sampling point. All vegetation collected along a transect was then combined for each species to constitute 1 sample. Vegetation was collected from a large enough number of in- dividuals so that individual plant variations in nutrient content would be minimized in the samples. Fall Fall samples were collected in early November, 1980. The 3 woody species collected for chemical analyses were taken along transects by the same method used to collect orange-hawkweed and sedge in the summer. Sample Preparation Summer Samples were placed in paper bags and dried at 60°C to a constant weight. Samples were reweighed after drying to determine percent moisture loss. Variation in moisture content of plants collected under varying climatic con- ditions was not taken into consideration. Researchers have determined that the nutritive values of various plant parts differ (Cook and Harris 1950, Dietz 1965, Bailey 1967). Cherry and jack pine leaves were separated from twigs after drying. By separating these 2 portions, values obtained through chemical analyses should represent the actual nutritive qualities of each plant part. 20 After parts were separated, samples were ground in a Wiley mill to pass a 1.0mm sieve. Ground samples were then stored in Whirl-paks (NASCO, Inc., Fort Atkinson, WI) for future chemical analyses. Fall Fall samples were handled differently than summer samples. During collection, all vegetation samples were clipped and placed in plastic bags. Samples were then frozen and stored until they could be dried and ground by methods stated previously. Chemical Analyses All vegetation samples were analyzed for percent dry matter, percent moisture, ash, crude protein (CP), ether extract (EE), ip ziggg dry matter digestibility (IVDMD), neutral-detergent fiber (NDF), acid-detergent fiber (ADF), acid-detergent lignin (ADL), and selected elements. Percent moisture and percent ash were determined by methods stated in A.O.A.C. (1975). Total nitrogen and phosphorus were determined with a Kjeldahl digestion method using a Tecator Block Digestor, Model DS-40 (Tecator, Inc , Boulder, CO). Once samples were digested, values were ob- tained using a Technicon Autoanalyzer II (Technicon Indus- trial Systems, Tarrytown, NY). Crude protein values were calculated as stated by A.O.A.C. (1975) using total 21 nitrogen values. Ether extract (crude fat) content was de- termined by methods stated in A.O.A.C. (1975) modified by weighing vegetation samples into tared filter paper ”packets” instead of thimbles. This modification enabled a larger number of samples to be analyzed per run. Percent EE was calculated as the weight loss in samples after extraction. Methods for NDF, ADF, and ADL analyses were those described by Goering and VanSoest (1970). 13 31533 dry matter di- gestibility was determined using the Tilley and Terry (1963) method, modified by the use of a phosphate-carbonate buffer solution to reduce foaming, and by reducing the amount of fluid used from 40ml to 10ml. Rumen fluid was obtained from a fistulated Holstein cow fed alfalfa hay and owned by Michigan State University's Department of Dairy Science. Percent dry matter analysis was conducted by weighing l.0-l.lg of dried, ground vegetation and putting it in aluminum pans which were then placed in an oven preheated to 100°C. Samples were dried for 24 hours, cooled in a dessicator, and then reweighed. Since each analysis dealt with only a small amount of sample, and all samples con- tained some moisture even after initial drying, it was imperative that the original sample weights be multiplied by percent dry matter. This calculation yielded the actual amount of vegetation used for an analysis. The percent of cell wall constituents (CWC) are the fiber contents (hemicellulose, cellulose, and lignin) 22 determined from NDF analysis. The cell soluble material (CSM) consisting of the soluble carbohydrates, starches, organic acids, protein and pectin was determined by sub- tracting CWC values from 100 (Goering and VanSoest 1970). The hemicellulose content of vegetation samples was calculated by subtracting ADF (cellulose and lignin) values from NDF (hemicellulose, cellulose, and lignin) values. Cellulose content was calculated by subtracting ADL (lignin) values from ADF content. Elemental analysis was conducted by the U.S. Forest Service - Michigan State University Cooperative Analytical Laboratory in East Lansing, Michigan. This analysis was conducted with a coupled plasma emission spectrophoto- meter (Spectrametrics, Inc., Andover, MA) (DeBolt 1980, Dahlquist and Knoll 1978). Statistical Analysis To determine if sludge treatment had a significant effect on the l) nutritional quality; 2) elemental content; and 3) annual productivity of vegetation collected during both sampling periods, a 1 way analysis of variance was performed on the data for all species. Bartlett's test was conducted on all data to test for homogenity of variance. Data which had heterogeneous variances were subjected to the arc sine transformation (Steel and Torrie 1980). Two- -way analysis of variance was used to determine the 23 season-treatment interaction of the 3 plant species collected during both seasons. A significance level of P i 0.10 was chosen to determine if significant differences between sludge-treated and control areas existed. Percent IVDMD data was used as reference to correlate fiber constituents and CSM values and IVDMD data. Cor- relations were made for all plant species on both sludge— treated and control areas to determine if the effects both treatments had on CSM and fiber constituent levels were related to IVDMD data. Regression equations were derived to determine if there was a relationship between height classes and annual productivity for cherries and jack pine. All regression equations were tested for significance using an F-test. Analysis of covariance was then used to determine if there was a significant difference in annual production on sludge- treated and control areas. RESULTS Annual Productivity The total annual productivity of 5 plant species <2m in height was 128% greater on sludge-treated areas than control areas. Cherries were the only species which dis- played a significant increase in production with sludge application (Table 3). The analysis of annual tree productivity by height showed a positive linear relationship between tree height and production (Table 4). Covariance analysis demonstrated that individual tree production was significantly greater on sludge—treated areas than controls for both plant species. Crude Protein Crude protein content was increased by sludge treatment in all plant species during both sampling periods (Table 5). Summer samples of jack pine twigs were the only species and part not to show significantly greater CP levels on sludge—treated areas. 25 Table 3. Above-ground net annual production (kg/ha) < 2 m height on study areas in 1980. Sludge- treated Control Species R SE X SE Significance Cherry 2084 185 867 369 P< 0.10 Jack pine 478 193 175 121 P> 0.10 Brambles 367 30 262 152 P3>O.lO Panic grass 168 61 51 12 P=>0.10 Other 11.6 3.3 6.5 1.6 P>’0-10 Total 3109 236 1362 210 PI<0.Ol 26 .mowooam zuon How AoH.ouvmv uchfiMflcme meB moaonm .modmaum>oo mo mwmhamcm Ou wcwpuooom mmmum Howucoo Eoum Amoco.ouvmv pamummwwp maucmowwmcwwm mm3 omwu Hmspw>wpcw mom coauoamowax 00. N Na xaa.m + o.sm- u A no. u Nu xoa.a + sum- N a« mafia acme mm. u my amm.a + m.NN- u a mma n ma me.N + cam- n as manage Houucoo mmummuuumwmdam mmwomam .ome CH A.aam maddwmv xwuoso mam Amcmwmxcmn mscwmv mama xomn mo AEUV uswwoc omwu pom ADB mun wv cowuonpoum Hmdccm masouwuw>0£m um: pom mwmzamcm mocmwum>oo mam mcowumsco Gowmwmuwom .q canny 27 omo.o w m mN.o No.m mm.o oo.m moanamnm omo.o w a wH.o om.m mN.o ma.o amaze «can xoam moo.o w a mm.o ao.m o~.o HH.HH mm>mma mama some ooa.c v a 53.0 mo.“ em.o wq.o meSD muumzo Hama moo.o w a om.o om.m mo.o aa.na mmapw oacmm moc.o w a m~.o w.o ma.o no.NH awemm moo.o w a Hm.o ea ma so.a o~.- emmsxsaa-mmcauo moo.c v m em.o 00 OH mm.o om.oa moHnEmwm coa.o M m ma.o mo.¢ mm.N mo.m mwfl3u mafia xomh omo.o w m oH.o cm.m mm.o ow.HH mo>moH mean xumm omo.o w a Na.o om.o mm.o wo.m awasu auumno ooa.o v m mm.o mo.ma Hm.m mw.mm mo>mmH xuwoso HmEEDm mm m mm M mumaflnmnoum Honucou mmumopuuowpaam momowmm moaned mHQEmm .ome .Hamm mam HoEEDm wcflhsm pmuomaaoo mmHaEmm pom mmmwm HouuCou mam poumopuumwpsam Eoww Cowumumwm> mo ucmucou ARV ammuowm opsuo mo mGOmfiwmmEoo .m mHan 28 Panic grass showed the greatest increase (133%) in CF content on sludge-treated areas during the summer. Analysis of fall samples showed brambles had the greatest response to sludge-treatment, increasing CP by 36%. All fall samples showed less variation in protein content be- tween treatments than did summer samples. Phosphorus Phosphorus content was consistently higher on sludge- treated areas than controls for all species during both sampling periods (Table 6). Significantly greater phos- phorus levels were found on sludge-treated areas than control areas for all species except cherry, jack pine, and bramble summer samples. Sedge and panic grass showed the greatest increase in phosphorus on sludge-treated areas. The phosphorus content was 140% and 126% higher on sludge-treated areas than controls for these species re- spectively. Ether Extract Ether extract content was not consistantly greater on sludge-treated areas than controls (Table 7). Panic grass was the only species to show significantly greater EE content on sludge-treated areas than controls. Ether extract content was 9% higher on sludge-treated areas. Orange-hawkweed displayed significantly greater values on control areas. 29 mood .V. 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Jack pine leaves and brambles showed signi- ficantly greater EE levels on control areas. Ash Analysis of summer samples showed only jack pine twigs had significantly greater ash content (an 11% increase) on sludge-treated areas than controls (Table 8). Those species which had significantly greater ash content on control areas were sedge and fall jack pine leaf samples. 1E vitro Dry Matter Digestibility IQ yipgg dry matter digestibility was significantly greater on sludge-treated areas than control areas for cherry leaves and twigs, brambles, and panic grass (Table 9). Panic grass showed the greatest increase in IVDMD (a 26% increase) on sludge-treated areas during the summer. For fall samples, brambles had the greatest in- crease (a 13% increase) in IVDMD on sludge-treated areas than control areas. Cell Wall Constituents Cell wall constituents were significantly lower on sludge-treated areas than controls for panic grass, and fall samples of jack pine twigs and brambles (Table 10). 32 ooa.o_Am no.0 Nq.m NH.o mq.m moanamum 2: .o M m 85 2A 85 24 mm?“ 21 x02. 035 v m 3.0 fia 85 cog 823: 32 x02. ooa.o Am ma.o mH.m HH.o mm.m mwazu xuuwno Hamm ooa.onwm NH.o «o.m no.0 mo.m mmmuw oflcmm 03.0; 2.0 $5. 3.0 £6 33% 03 .o A a 3.0 13.0 and $20 cmmsx3m55wcmuo ooa.c.mm NH.o mm.m mN.o oo.m moanamum 03.0 v a 8.0 2: Ed Ea Amps .23 x02. 2:5 A m 8.0 34 3.0 a; 8%3 33 x22. ooH.o_Am mH.o om.m oa.o om.N me3u anyono 2: .o A m 2.0 2 .m S .o 33 85.3 3.85 umEESm mm m mm m mufiHwnmnopm Houucou vmumouuumwpsam mmflommm powpmm mHaEmm .omma .HHmm paw uoaesm wcwusp pmuooaaoo monEwm pom mmoum Houucoo cam pmummuuuowpsHm boum cowumummm> mo ucmucoo Axv 5mm Mo mGOmwquEoo .m maame 33 ooa.ou.m mo.a No.5N o©.o No.om mmanamum ooH.o_Am so.o NH.¢m mm.o mm.mm mafia“ mafia xomw ooa.ohwm Hq.o ¢o.Hm mm.o qu.am mm>moH mafia xomh ooH.ouvm Ha.o HA.©N mo.H No.0N awasu suumso Hamm oHo.o Vm em.m ma.mm mm.o mm.aa mmmuw uacmm ooa.o_Am mm.~ NN.mm ¢N.m om.am wwwmm ooH.0nwm HN.N am.¢m om.H on.qm cmmsxsms-mmamuo cmo.o Va mN.H mm.qm NH.H mm.oq mmHQEmpm ooa.o.Am mm.o om.mm ca.H am as mmasu mafia xomn ooa.ohmm No.H ma.mm om.o om am mm>mmH mafia acme omo.o.wm mo.o mm.m~ «w.o mo mm mwasu suumno coa.o vm mo.m m .n¢ wo.m om.om mm>mma muumsu “mafidm mm I mm m kuHHHnmnoum Hopucoo woumouunmwwsam mmwommm cowhmm maaamm .ome .Hamm paw HmEEDm quudn pouomaaoo mmHaEmm pom wmoum Houucoo Umm pmummuuumwwsam Eonm cowumuoum> wo va xuwaflnaumowflw umuwe hub onuw> Cw mo maomwumaaoo O .o manme 34 0m0.ouvm 00.0 no.5m mm.N0 m~.0 0m.0q 0n.mm moamamum w “Bu mN0.0UVm 00.H oq.m0 00.0¢ m0.H 00.0w NH.mq ocfia xomh mm>mmH 2:5 v m 2.0 3.8 3.3 24 3.8 Ram 23 x02. 00H.0uvm 05.0 ¢H.0m 00.m0 0H.H wq.wm mm.H0 mwflzu muumso Hamm m00.0mnm mm.0 N0.NN 00.mm MH.H H¢.0N mm.am mmmhw owamm 2:5 v m $5 3.3 3.: 8.0 $33 :50 mwumm 0mm3x3mn 00H.0Uvm “0.0 H0.q0 mm.mm Hm.H Nm.m0 00.¢m umwcmuo 00H.0uvm 0N.H HH.nm 00.Nq m¢.a 00.mm «0.qq moanamum mwfl3u 00H.0uvm 00.0 Hm.m¢ 00.Qm 0H.H ¢0.n¢ 00.mm mafia xowh m mo>mmH 00H.0U.m 0¢.N qfi.0¢ 00.0w 00.H qq.¢m 0m.mq mafia xomh 00H.0“vm 0m.N mm.qm m0.m0 No.0 mm.mm mm.oo wwwzu zuumso mo>moH 00H.0U.m 0H.m 0m.wo H<.Hm qq.m mm.Hu mq.wm ~AHuono uoEEDm Emu 030 Emu 030 3332on mm m mm m moflooam pOHHmQ onEmm .000H .Hamw paw HwEEsm wcflunp vmuooaaoo mmamemm you mwmum Houucoo paw pmumouusmprHm Eoum coauMumwm> mo ucoucoo ANEmuv Hmwumume mHQDHOmuHHoo paw Aflozov mudmfiuwumcoo Ham3uHHmo mo accommodaoo .0H oHan 35 Cell Soluble Material Summer samples of panic grass and fall samples of jack pine twigs and brambles showed significantly greater CSM contents on sludge-treated areas than control areas (Table 10). Acid-Detergent Fiber Cellulose and lignin content of vegetation, de- termined by the ADF analysis was consistantly lower on sludge-treated areas than controls for all plant species (Table 11). This general trend was observed in vegetation samples for both sampling periods. Significantly lower values were found in jack pine twigs, brambles, orange- hawkweed, sedge, and panic grass for summer samples. Brambles were also significantly lower in ADF on sludge— treated areas than controls during the fall. Orange-hawkweed and brambles showed the greatest change in ADF content with sludge application for the summer and fall respectively. Acid-detergent fiber values for orange-hawkweed and fall bramble samples were 15% and 7% lower on sludge-treated areas respectively. Hemicellulose Fall jack pine twig samples were the only species and part to show significantly lower hemicellulose content on sludge-treated areas than control areas. Orange-hawkweed 36 ooa.o..m mm H «N mm Hm.o Na we mmansmum ooa.o_Am om.H a¢.am ma.o HN.om mwazu mafia xumw ooH.o.Am mo.o Ho.om a~.o oq.m~ mm>mmH «can xomw ooa.o_Am SH.H wa.mm ww.o aq.~m mwa3u suumnu Hamm moo.0mHm Hm.o oo.mq am.o mm.mm Ammum uacmm ooa.omwm oo.c mm.mq am.o mm.oq mmemm moo.omwm ma.o No.mm mn.o ao.m~ umosxzmn-mwcmuo mmo.o.wm H¢.H wo.mm H¢.H mo.~m mmHnamum ooH.o vm mm.o m .oq no.0 Ho.q¢ maps“ «can xomn ooH.o_Am mN.m mo.aq Ho.H om.om mm>mmH mafia xumw ooH.c Am ww.o 0N.om ma.o HH.on mwazu auumnu ooa.o.Am Hm.N mm.q~ 0H.H aq.oH mm>mmH >Humno .HmEESm mm M mm m muflawnmnoum Houucoo woummuuumwpsam mmfloomm powhma mHmEmm .owma .HHmw paw HmEESm unwuss Umuooaaoo mmamamm How mmmuw Houucoo paw pmumouu ummpdam Scum coaumummm> mo ucoucoo ANV umnwm ucomumumpupflom mo mCOmemano .HH mafima 37 and bramble (summer and fall) samples had significantly greater hemicellulose values on control areas than sludge-treated sites (Table 12). Acid-Detergent Lignin Acid-detergent lignin content was consistantly lower on sludge—treated areas than controls for all species except sedge (Table 13). Summer species and parts that showed significantly lower ADL content on sludge-treated areas were: jack pine twigs, brambles, and cherry twigs. Brambles also showed significantly lower ADL content on sludge-treated areas than control areas during the fall. This species had the greatest decrease in ADL content with sludge-treatment for summer and fall (48% and 10% re- spectively). Cellulose All herbaceous plant species collected showed signi- ficantly lower cellulose content on sludge-treated areas than control areas (Table 14). Both panic grass and orange-hawkweed showed the greatest change in cellulose content with sludge application. Cellulose values were 16% lower on sludge-treated areas for both species. Elemental Analysis The results of statistical analysis of element con- tents in plant samples for sludge-treated and control 38 moo.om”m mm.o Nq.a mm.o mm.oa mmanamum ooH.o.vm mo.H HH.m am.o om.n mwazu mafia xomw ooH.o_Am mm.o mm.o om.H em.“ mm>mmH mean some ooa.o_Am ao.a ao.m No.0 mo.m Amazu muumno Hana ooH.o_Am am.o aq.am «0.0 mo.am mmmuw owcmm ooH.o mm mm.o mm.m~ HH.H mH.mN «wumm mN0.0.wm 0m.0 mm.m mw.0 H¢.m pmm3x3mnumwcmno moo.o..m om.o Hm. wa.o H¢.HH . mmHQEmum ooa.o_Am om.o am.a mm.o om.e Amazu «can xomw ooa.o;Am mo.m o~.~a aq.H om.m mm>mmH mafia xumn ooH.c Am ON.N Na.m qa.o «H.0H Amazu Apumno QQH.c Am ma.H mw.w H¢.N mm.w mm>mmH suumno Hoaanm mm M mm M. muHHflnmpoum HOpucoo pmummuunowpaam mmwomam poauoa mHQEmm .ome .Hamm paw nmEESw waHSp pmuomaaoo mmaafimm now mmmhm Honucoo paw cmummuunmmpsam Ecum Cowumuoum> mo unmucoo ARV mmoHDHHmoHEmn mo mGOmHHMQEoo .NH mHLmH 39 ooa.o..m Ho.o Hm.©H oN.o mm.mH mmagamum coa.c.Am Hm.o am.ma No.0 ow.aa mafia“ mafia xomw coa.o.Am Hq.o «w.NH om.o am.~a mm>mmH mane acme coH.o.Am aw.H ew.mN oo.H mm.a~ mwasu Anyone Hana oca.o_Am NH.o ow.m wH.o Ho.m mmmuw uwcmm coa.o_Am ~@.o oa.m ma.o om.“ wwwmm ooa. .Mafl aq.o NH.o mo.o mq.m wmmzxzmn-mwampo oca.omHm wq.a oo.HH mm.o mm.a mmanamum moo.ouvm oo.o mo.- mm.o mm.HN mwasu mama xomn ooa.o mm mm.o mm.oH co.H mm.ma mm>mmH mafia xumm ooH.o Va“ am.H wa.om mw.o NH.HN wwwzu suuwnu ooa.o.Am SH.H aN.HH Nw.o ma.a mm>mmH supmnu .Hmfiafim mm m mm m muflHmeQOHm Houucoo poummuuumwpsam mmwommm powwow mHmEmm .OmmH .HHmm USN .HmeEDm wCHHDmV UmuomHHOU mmHQEmm MOM mwmhw HOHHCOU Ufim fimummub nompsam Eonm cowumumwm> mo ucoucoo ARV cflcwwa ucwmuouwpnpwom mo wCOmHHmQEou .mH manme 40 000.0.00 00.0 00.00 00.0 00 00 00005000 000.0_A0 00.0 00 00 00.0 00.00 00030 0000 0000 000.0 A0 00.0 00.00 00.0 00.00 003000 0000 0000 000.0.A0 00.0 00.00 00.0 00.00 00030 000000 0000 000.0000 00.0 00.00 00.0 00.00 00000 00000 000.0.00 00.0 00.00 00.0 00.00 00000 000.0“.0 00.0 00.00 00.0 00.00 00030300-000000 000.0_A0 00.0 00.00 00.0 00.00 00005000 000.0 A0 00.0 00.00 00.0 00.00 00030 0:00 0000 000.0_A0 00.0 00.00 00.0 00 00 003000 0000 0000 000.0_A0 00.0 00.00 00.0 00.00 00030 000000 000.0 A0 00.0 00.00 00.0 00.00 003000 000000 005550 00 m 00 m mu0awnmnoum HOpucou pmumouuumeSHm mm0omam 0000mm mHQEmm .0000 .Hamw 0cm umbasm uCHHDp vmuowHHoo mmamfimm How mmmhm Houucoo paw poummuuummpsam Eoum EOHumumwm> mo ucoucoo ANV mmoHSHHmo mo 0com0umquo .qH manme 41 areas collected in 1980 are presented in Tables 15 through 27. Panic grass and brambles (summer and fall) assimi- lated significantly higher concentrations for the most elements on sludge-treated areas. Statistical analysis of summer data showed woody species and parts (cherries, jack pine, and brambles) accumulated significantly greater metal concentrations on sludge-treated areas than controls for an average of 2 elements per species and part. Orange- hawkweed had significantly greater concentrations for 4 of the selected metals on sludge—treated areas. Potassium was the only element significantly greater in sedge on sludge-treated sites. Panic grass accumulated the most elements during the summer. This species showed signifi- cantly greater concentrations for 11 of the 13 elements on sludge-treated areas than controls. Analysis of fall samples indicated that brambles assimilated the most elements on sludge-treated sites. From summer to fall this species accumulated significantly greater concentrations of 6 additional elements on sludge- treated areas than controls. Cadmium was the only element with significantly greater concentrations on sludge-treated areas than controls in cherry twig and jack pine leaf samples. 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Ucm mmummuuummwdam Eoum coaumumwm>.mfiu ucmucbo dfiv Efifluamo mo wCOmwummEou .NN manme 55 Season-Treatment Interactions Significant interactions between seasons and treatment effects on all chemical analyses are shown in Table 28. The magnitude of the effects of sludge application and season varied among the 4 plant species sampled during both seasons. Brambles showed significant interactions between seasons and treatments for the most elements and nutrients in which analyses were conducted. Tn Vitro Dry Matter Digestibility-Fiber Constituents and Cell Soluble Material Correlations Correlations for plant species on sludge-treated areas showed there was not a close relationship between all fiber constituents and CSM and IVDMD for any 1 species (Table 29). A strong correlation, however, was determined in jack pine twigs for hemicellulose and lignin to IVDMD. Orange- hawkweed also showed a strong relationship between IVDMD and cellulose and CSM. A strong relationship was also determined in sedge for CSM as well as lignin content Correlation for fall cherry twig samples showed a strong relationship between all fiber constituents and IVDMD, however, not with CSM. 56 .oa.o um mooConmmmwo unmoHMflcmHm monsoon»,n wooq.mm «seq.m ammo.m amok.ma aqao.w awmm.m «mmm.NN moanamum £25 Names $2.0 $25 amps mafia xomh aooo.m *woa.m mo>mmH mafia 30mm «935 mwafi %Hno£o mm< mm Esfiwmmuom Eoflmmcwmz EDSwEDH< ommA Sbwfionzo meowz afifloamu CouH mandamus: pom mquEmHm mmfiomam .mmaafimm coaumumwo> mo mucoucoo udowuusc mam ucoEoHo mo chmmom wow mucoEumonu CmoBuoa msofluomumucw oomUHMflcme mo moDHm>um .wN oHan 57 HmHHmumE mannaom Hamo p 5505550 omoHSHHmon m 50.0 50 55005555 5005 mmoHDHHoowammm no.0 um unmoHMchHm « 55.0 0050.0- 00.0- 00.0 05.0 05.0 50.0- 05.0 00505050 mw53u 00.0 50.0 05.0- 05.0 00.0 00.0- 00.0 00.0- 0:50 5000 mo>moH 50.0 05.0- 00.0- 50.0- 55.0 05.0 005.0- 05.0- 0550 5000 05.0 «000.0- 0000.0 000.0 00.0 50.0- 00.0- 55.0 00535 555000 uuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuu 55mm nulls-uxuuuun-unun-nunun-annunnn- 00.0 05.0- 05.0- 00.0 55 0- 05.0 05.0 50.0 00050 05505 000.0 000.0- 05.0- 05.0- 50.0 05.0- 00.0 05.0- 00000 omm3x3m£ 505.0 50.0- 50.0- 50.0- 050.0- 50.0 00.0 «550.0 -005050 00.0- 05.0 50.0- 50.0- 00.0- 55.0- 00.0 55.0- 00505050 mw53u 50.0 00.0- 05.0 05.0 00.0- 0000.0- 05.0 005.0 0550 5000 mm>mo5 05.0- 50.0- 00.0- 00.0 05.0 00.0 05.0 55.0- 0:50 5000 00.0- 00.0- 50.0 50.0 50.0- 50.0 00.0- 00.0- 00535 555000 005.0 00.0- 0050.0- 00.0- 05.0- 05.0- 05.0- 50.0 002005 555000 uuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuu HoEEDm n-uun-u-nnun-uuu-sununununuuuu-u :00 0055 5500 5500 0200 05555 05500 05900 5055500 ooummuu-mwodam mmwomam .Hm55mumm manaaomuaaoo cam muconufiumaoo Honwm paw %555525500w5o Hmuume who 0555> 55 Cmmsumn mufim5owmmooo cowuwamuuoo maaawm .mm manme DISCUSSION Annual Productivity The application of additional nutrients by inorganic and organic fertilizers to agricultural crops and forest lands has been documented by many authors to increase annual productivity (Sopper and Kardos 1973, Leaf et al. 1975, Bayoumi and Smith 1976, Hinesly gt El- 1979). The significant increase in total annual productivity (Table 3) which occurred in this study as a result of sludge application parallels what these other researchers found. Hahn at al. (unpubl. data) notes that spray irrigating, anaerobically digested, domestic sludge to pasture grasses significantly increased yields. Sopper and Kardos (1973) observed a similar response of increased annual production of an old field irrigated with municipal wastewater. They noted that with irrigation, productivity increased an average of 201% annually over a lO-year period. Red pine (Pinus resinosa) plots were also irrigated with waste- water in Sopper and Kardos' (1973) project and annual productivity monitored. They noted that with irrigation both the diameter and height of red pine increased. 58 59 Results from these projects support the findings of this study, that the application of organic fertilizers in- creased the productivity of numerous vegetation species. The positive responses of various vegetation species to organic fertilizers has also been produced with in- organic fertilizers (Thomas gt al. 1963, Bayoumi and Smith 1976). These authors documented that the application of nitrogen fertilizers increased forage production of both herbaceous and woody species. The response of pin cherries to fertilization and heavy cutting has been documented by Auchmoody (1979). He stated that pin cherries regenerate heavily after a disturbance, such as cutting or fire but that many seeds may remain dormant in the soil for years. Auchmoody (1979) noted, however, seeds may be stimulated to germinate with an application of nitrogen as low as 56 kg/ha. Because the addressed study area was clear-cut and then treated with sludge which contained the equivalent of 481.5 kg/ha of nitrogen this would be expected to stimulate cherry germination and production. The clear-cutting in combi- nation with sludge application would thus cause signifi- cantly greater annual productivity on sludge-treated areas. Another factor which may have contributed to the signi- ficant increase in pin cherry production is the high annual productivity and nutrient accumulation pin cherry normally exhibits (Marks 1974, Covington and Aber 1980). 6O Safford and Filip (1974) noted that with the ferti- lization of pin cherry the proportion and absolute amount of nutrients shifted from an even distribution among species in an unfertilized stand to a high proportion of pin cherry. They also documented that with fertilization 4 years after clear-cutting, Bgtgg spp. as well as pin cherries tended to dominate the stand in above-ground bio- mass and number of seedlings. Woodyard (1982) noted this same response with sludge application to the jack pine clear-cut of this study. Since sludge application did increase production, it is possible that the initial effects of application are advantageous to wildlife (increased quantity of vegetation) but the long term effects may be detrimental. Because the nutrients contained in sludge are accelerating the pro- duction of plants they are also contributing to accelerating the maturation of those plants. Therefore, although plants fertilized with sludge may provide more quality vegetation for food and cover initially, the amount of time these areas will be beneficial to wildlife may be short-lived as vegetation structure changes. The preliminary findings of West gt gl. (1981) who monitored small mammal and herbivore use of sludge-treated areas support this hypo- thesis. 61 Crude Protein From the data obtained it is evident that a single application of non-industrial municipal sludge did im- prove the nutritional quality of most wildlife forage species. The degree of nutritive quality improvement varied by plant species and season. Because protein is a principle component of an animal's soft structures, a constant and adequate amount is re- quired in the diet to support the animal's growth and reproduction (Maynard gt gt. 1979). The protein requirements of an animal, however, will vary with species and age. Murphy and Coates (1966) documented that forage protein content may be the most critical nutrient on some ranges and may account for low production and physical development observed in white-tailed deer (Odocoileus virginianus). They noted that body weights and antler development of yearlings and adult male white-tailed deer were drastically retarded by a diet of 7% protein. This same effect on white-tailed deer subsisting on protein-poor forages has been noted by other researchers (Einarsen 1946, French gt gt. 1956, McEwen gt gt. 1957, Smith gt gt. 1975). Because of the nitrogen contained in sludge and its' application in this study to an early successional site with nutrient—poor soil, the nitrogen was readily assi- milated by the vegetation. With this nitrogen influx, none of the plant species on sludge-treated areas had CP 62 levels below the minimum requirement of 7% required by white-tailed deer. However, on control areas, 42% of the plant species of both seasons exhibited CP levels less than 7%. Because of the sludge-induced increase in GP content and production of all plant species, wildlife species, such as white—tailed deer may choose to use sludge-treated vegetation more heavily than unsludged. Sludge-treated areas will thus provide more cover and better quality of forage for wildlife. Woodyard (1982) documented that the percentage of browsed stems for cherries and brambles was significantly greater (P <0 01) on sludge-treated areas than controls on this study site. These results suggest that wildlife use was greater on sludge-treated areas possibly because of the increase in the quantity and quality of vegetation. Results from other studies support the finding that wildlife species do select areas where plant quantity and quality are increased with fertilization (Leaf gt gt. 1975, Dressler and Wood 1976, Barrett 1979). Since cherries are known to accumulate nutrients readily and retain these primarily in the leaves and twigs (Safford and Filip 1974), these species showed greater CP levels than other plant species during both seasons. This corresponds with cherries' ability to assimilate high ni- trogen levels. 63 It has been documented by Safford and Filip (1974) that figtgg spp. may also (like pin cherry) accumulate nu- trients readily. This ability may enable ggtgg spp. to dominate a stand in total biomass after fertilization. This was evident since brambles showed a greater increase in GP content on sludge-treated areas than control areas in the fall. Phosphorus PhOSphorus has 3 basic functions in an organism; l) structural,2) a central role in energy transformation, and 3) a component of the nucleoproteins and cytoplasm in cells (Maynard gt gt. 1979). The ability of an organism to meet its phosphorus requirements will depend on the amount contained in the soil available for plant assimilation. The application of 571.4 kg/ha of phosphorus from sludge application provided plants with additional nutrients for plant accumulation. The amount of phosphorus accumulated by vegetation, however, may have resulted from: 1) the phosphorus content of the sludge, 2) the amount and chemical form of phosphorus available on the site prior to sludge application, 3) the speed at which different plant species grow and assimilate nutrients, and 4) root depths of various species. As stated by Sommers (1977a), phosphorus concentrations in sewage sludge may equal or exceed the amount of nitrogen 6A present. Because the phosphorus content in the sludge was slightly higher than the total nitrogen, the phosphorus content of this sludge was considered average. Some plant species accumulated significantly greater concentrations on sludge-treated areas than control areas. Brockway gt gt. (1979) noted that irrigating vegetation with wastewater caused an increase in the level of phos- phorus in vegetation. However, he attributed this increase to applying wastewater to a site already low in phosphorus. As stated earlier, the soil on the site was a nutrient- deficient sandy soil which may have been the reason vegetation accumulated phosphorus so readily. From the results of summer analyses, only the herbaceous species accumulated significantly greater concentrations of phosphorus on sludge-treated areas than controls (Table 6). This response may be attributed to herbaceous species having a shallower root system than woody species and thus being capable of assimilating nutrients in the surface soil layers faster. Because the nutrients contained in the sludge will only enter the surface soil layers by de- composition and leaching resulting from percolating rain- fall (Brockway 1979), the movement of phosphorus may be limited by weather and climate. As shown in Figure 2, the amount of precipitation during the summer sampling period (mid-July to mid-August) was well below average. This lack of rainfall may have contributed to the low 65 mobility of phosphorus. Because phosphorus was restricted to the upper soil layers, only species with shallow root systems (such as herbaceous species) could assimilate sig- nificant levels. Analysis of fall samples showed that all woody species had significantly greater phosphorus levels on sludge- treated areas than controls (Table 6). This positive re- sponse of phosphorus accumulation in the fall and not in summer may also be attributed to the amount of rainfall. Figure 2 shows that precipitation during the fall (September through November) was much greater than during the summer. This increase in precipitation may have caused phosphorus to be leached into deeper soil layers making it available to plants with a deep root system (such as woody species). As documented by Short gt gt. (1966), the phosphorus content in a variety of vegetation samples has been re- lated to protein content. This same type of response was observed in this study with most species. Those plant species which showed a significant increase in phosphorus, as a result of sludge-treatment, also showed an increase in CF (Table 5). However, all species which displayed a significant increase in GP did not show a significant rise in phosphorus. This response could be attributed to the sludge containing a relatively high concentration of ni- trogen in relation to phosphorus (Sopper and Kardos 1973). 66 Because of the important functions phosphorus plays in an organism, adequate dietary supplies are essential. As discussed, sludge application did increase phosphorus content of a few plant species; was sludge application, however, necessary to increase the phosphorus content of vegetation to meet dietary needs of wildlife species? Ullrey (pers. comm.) stated that weaned white-tailed deer fawns require a minimum of 0.28% (2800ppm — dry rratter basis) phosphorus when the diet contains 0.45% (4500ppm) calcium for growth, skeletal, and antler development (calcium phosphorus ratios will be discussed later). The results of this study indicate that only 25% of the summer plant species and parts on control areas met these re- quirements. Sludge-treated areas, however, showed adequate phosphorus levels for 63% of the plant species and parts. None of the fall plant species on sludge-treated or control areas had phosphorus levels of at least 0.28%. These results indicate that sludge application may increase normally low phosphorus levels to meet the dietary require- ments of wildlife species, such as the white-tailed deer. Ether Extract Fats in the diet of an animal are essential for: 1) providing 2.25 times the metabolizable energy as proteins and carbohydrates (also serve as an energy source to plants), 2) aiding in the palatability of forage, 3) aiding in 67 carrying the fat solublexfixandns,andl0 supplying essential fatty acids. Because of these factors, EE is an important parameter to assess when evaluating the nutritional quality of forages. In interpreting EB data, however, it should be realized that ether, in addition to extracting lipids also extracts chlorophyll, xanthophyll, carotene, waxes, and other substances. Therefore, the EB content of forages should not be referred to solely as fat (Maynard _t _t. 1979). As the results of this project indicate, EE content was not consistantly greater on either sludge-treated or control areas. This inconsistancy may have been pro- duced by the level of l nutrient affecting the relative amount of another nutrient in a plant. If a plant has a higher relative amount of other chemical constituents (nitrogen and phosphorus) then the relative amount of EB will decline without an actual change in amount. There- fore, the relative amounts of EB would not be expected to increase with sludge application because of the accumulation of nitrogen and phosphorus into the cell contents. Results of the chemical analysis support this conclusion since species on control areas showed a trend of higher EE content. Ash The ash content of vegetation samples comprised very little of the total composition ( 0.5 ppm did not have deficient levels (based on the dietary requirements of sheep) of the above mentioned elements or protein (National Research Council 1968), cadmium toxicity problems may not occur. The subject of heavy metal accumulation in animal tissues is the focus of ongoing research. Magnesium, like cadmium, also exhibited concentrations which exceed the maximum tolerance level (0.5-0.08%) (National Research Council 1980) on both sludge-treated and control areas (signi- ficantly greater on sludge-treated areas). As documented by the National Research Council (1980), however, toxicosis due to ingestion of natural forages has not been reported and does not appear likely. This may be attributed to adequate calcium—phosphorus ratios (1.5 1) which protect an animal from toxicosis. Because most plant species on sludge- treated areas (jack pine leaves - summer and fall, jack pine twigs fall, and sedge on control areas) had calcium- phosphorus ratios between 2:1 - 1:1, as mentioned earlier, magnesium toxicosis is unlikely. Treatment-Season Interactions The significance of conducting this analysis was to test if the response of different chemical constituents and elements to sludge application was significantly different between seasons. Thus, significant results would indicate 77 that both factors (treatments and seasons) are dependent on one another in influencing nutrient and element levels. Brambles showed significant interactions between treat- ments and seasons for 7 of the chemical analyses conducted (Table 28). Other species had less of a response, thus the effects of treatments and seasons must act more inde- pendently in affecting the nutrient quality of these species. These significant interactions between treatments and seasons indicate that there are various combinations from sludge-treatement and seasons which may influence the nutritive quality of vegetation. SUMMARY AND RECOMMENDATIONS The application of non-industrial municipal sludge in- creased plant productivity and the nutrient quality of a variety of wildlife forage species. These results may be attributed to the influx of nutrients into a nutrient poor forest soil. The results of this study show an application of non- industrial municipal sludge, prior to the growing season, may increase the quantity and quality of vegetation for wildlife use. Plant species which will benefit the most from additional nutrients readily are species which charac- teristically assimilate nutrients readily and/or those which have shallow root systems. These species will benefit primarily because sludge-borne nutrients may be restricted to surface soil layers. The assimilation of these nutrients by plants will aid in maintaining wildlife forage species in a productive state of vigor which is attractive to wildlife. Because of the benefits received by applying sludge to appropriate forest lands, sludge application may serve as a method of habitat improvement for a diversity of wild- life species. 79 The emphasis of this study was to identify the potential of sludge application to forested areas as a disposal al- ternative and method of habitat improvement. Because sludge application increased production as well as nutrient quality, plant maturation may be accelerated and therefore, the benefits of sludge application short-lived. Infor- mation regarding the duration of sludge application benefits F to various vegetation types and the accumulation of heavy metals in both forages and animal tissues should be the focus of future research. This will provide wildlife 5 managers important data enabling them to maintain desirable forage species in a high nutritional status by sludge-treating various areas on a rotational basis. LITERATURE CITED LITERATURE CITED AOAC. 1975. Offical methods of analysis of the association of official analytical chemists. 12th ed. Assoc. Off. Anal. Chem. 1094 pp. Auchmoody, L. R. 1979. Nitrogen fertilization stimulates germination of dormant pin cherry seed. Can J. For. Res. 9:514-516. Bailey, J. A. 1967. Sampling deer browse for crude protein. J. Wildl. Manage. 31 437-442. Bailey, J. A. 1968. Effects of soil fertilization on the concentration of crude protein in witchhobble browse. N.Y. Fish and Game J. 15:155-164. Barrett, M. W. 1979. 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Wildlife response to sludge ap- plication. M.S. Thesis. Michigan State University, East Lansing. 64 pp. APPENDIX Table 30. 87 List of vascular plants on the study site. Common name Scientific name Family Jack pine ap Pinus banksiana, Lamb Pinaceae Bigtooth aspen Populus grandidentata, Salicaceae Michx. Quaking aspen Populus tremuloides, Salicaceae Michx. Red cedar Juniperus virginiana, Cupressaceae L. White spruce Picea glauca, Pinaceae (Moench) Voss. Black cherry Prunus serotina, Ehrh. Rosaceae Choke cherry (ap) Prunus virginiana, L. Roseceae Pin cherry Prunus pennsylvanica, Rosaceae L. Red maple Acer rubrum, L. Aceraceae American red Rubus strigosus, Rosaceae raspberry (ap) Michx. Common blackberry Rubus allegheniensis, Rosaceae Porter. Cinquefoil Potentilla recta, L. Rosaceae Mapleleaf Viburnum Viburnum acerfolium, Caprifoliaceae L. Panic grass bp Panicum virgatum, L. Gramineae Sedge b Carex spp., L. Cyperaceae Red sorrel Rumex Acetosella, L. Polygonaceae Smartweed Polygonum spp., L. Polygonaceae Goldenrod Solidago spp., L. Compositae Spotted knapweed Centaurea maculosa, Compositae Lam. Bull thistle Cirsium vulgare, Compositae (Savi) Tenore. Pussytoes Antennaria neglecta, Compositae Green. Orange-hawkweed b Hieracium aurantiacum, Compositae L. Aster Aster spp., L. Compositae 88 Table 3M1 (cont'd.) Common name Scientific name Family Pokeweed Phytolacca americana, Phytolaccaceae L. Tomato Lyc0persicon Solanaceae esculentum, Mill. Common mullein Verbascum Thapsus, L. Scrophulariaceae Common plantain Plantago major, L. Plantaginaceae St. John's-wort Hypericum perforatum, Hypericaceae L. adenotes collection during the summer and fall bdenotes collection during just summer pspecies collected for annual productivity analysis "7'111111111171111111115