m‘fi—w— FRiMARY PRGEEU‘CTIGN, ENERGETECS. ANEE NUTRiENT UTIUZATION IN A WARM..WA?ER STREAM Thesis ‘or {he Degree 05 pk. D. MICHIGAN STATE UNEVERSITY Alfred Richard Grzenda 1.960 This is to certify that the thesis entitled‘ PRIMARY PRODUCTION, ENERGETICS, AND NUTRIENT UTILIZATION IN A WARM-WATER STREAM presented by Alfred Richard Grzenda has been accepted towards fulfillment of the requirements for Ph. D degree in Fisheries and Wildlife \<\> thkt Q\ @CKQQ Robert C. Bal 1 Major professor Date May 2, 1960 0-169 LIBRARY Michigan State University 7 "yank-J: . 94",: K~1I".G~--'w' OVERDUE FINES: 25¢ per day per item RETURNING LIBRARY MATERIALS: Place in book return to remove charge from circulation records PRIMARY PRODUCTION, ENERGETICS, AND NUTRIENT UTILIZATION IN A wmi-WATER STREAM by ALFRED RICHARD GRZENDA AN ABSTRACT Suhnitted to the School for Advanced Graduate Studies of Michigan State University of Agriculture and Applied Science in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Fisheries and Wildlife 1960 - m that G Rafi . ‘\ l | l \ Alfred Richard Grzenda Primary production, energetics, and nutrient fixation were investi- gated in a warm-water stream located in the south central portion of the Lower Pennisula of Michigan. Gravimetric net production estimates were made from periphyton har- vested from plexiglass substrata which had been sumerged in the stream. Colorimetric productivity measurements were made by relating the absor- bancy of phytOpigments extracted from the periphyton colony to the ash- free dry weight of the colony. Good agreement was noted between the two methods. Diurnal oxygen curves were found to be unsuitable for year- round productivity measurements. Seasonal production'maxima.were observed in late spring, early summer, and late fall. The seasonal range in net primary production as measured by artificial substrata is 0.01 to 2.28 -2 2 m day-1. The annual mean rate of gross production is estimated to be about 1 g m-2 day-1. g day-1 with an annual mean of 0.56 g m- A model describing primary energetics was synthesized from observed data and literature values. Photosynthetic efficiencies based on net production and surface radiation varied from 0.003 to 0.2h5% with an annual mean of 0.07%. The annual mean efficiency based on gross produc- tion and surface energy was estimated. to be 0.1%. The annual phosphorus import into the study area was 16 metric tons. During three flood periods collectively representing a time lapse of 30 days about hS% of the annual import was carried into the study area. The total inorganic nitrogen import for a nine month period was 8h.metric tons. The effect of floods on the seasonal distribution of inorganic nitrogen was less pronounced than in the case of phosphorus. The data indicate that sanitary drains are the major source of phosphorus and surface run- off the major source of inorganic nitrogen. The phosphorus content of periphyton varied considerably throughout the year. In genera1,the slowest growing colonies had the highest phos- phorus content. The ratio of phosphorus content to weight of the peri- phyton colony (x 1,000) varied from 1.Sh to 33.63. During periods of maximal periphyton production the ratio of phosphorus to nitrogen within colony was approximately 1. Seasonal phosphorus utilization efficiency quotients based on sol- uble phosphorus import and calculated for a 100 meter length of stream varied from 0.0007 to 0.61% depending upon the growth rate of the peri- phyton. The lowest efficiencies were observed during the mid-winter production minima.. Similar quotients computed for inorganic nitrogen ranged from 0.03 to 0.19%. The ratio of phytopigment absorbancy to periphyton colony weight can be used to differentiate between oligosaprobic and mesosaprobic communities. Such ratios obtained from a clean-water zone vary from 10.0 to 16.9. Similar values obtained from a polluted zone vary from 1.8 to 2.9. Ratios based on the phosphorus content of the colony are less reli- able indices of pollution and appear to be limited to pollution detection rather than evaluation. PHILARY PRODUCTION, ENERGETICS, AND NUTRIENT UTILIZATION IN A WARE-WATER STREAM by ALFRED RICHARD GRZENDA A THESIS Submitted to the School for Advanced Graduate Studies of iichigan State University of Agriculture and Applied Science in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Fisheries and Wildlife 1960 (D 2. :93 ',: 4'. ii f. I . . ‘v ' 4 Q {in} “I .I I; "‘1‘" t I wish to extend my sincere appreciation and indebtedness to Dr. Robert 0. Ball for supervising this study. His comments concerning the field work and the preparation of this manuscript were invaluable. I wish to thank Drs. Gordon Guyer, Gerald w. Prescott, and Peter I. Tack for the time they spent serving as members of my guidance committee. I am also grateful to Dr.‘£orris L. Brehmer andlir. David L. Correll for their advice and assistance in conducting laboratory analyses. I wish to extend my sincere thanks for the generous financial assist- ance given to me by the Department of Fisheries and Wildlife, Nichigan State University; and the Division of Research Grants of the National Institutes of Health, Deparbnent of Health, Education, and welfare. TABLE OF CONTENTS INTRODUCTIOIJ 00.000.000.000...OOOOOOOOOOOOO0.0.0000000000000000 DESCRIPTION OF STUDY AREA . . . . . .............................. . r‘xmoos mm EQUIHI‘LENT ................ . ..... . pH and alkalinity .................................... Turbidity ............................................ Seston ............................................... Absorbancy ........................................... Phosphorus ........................................... Nitrogen ............................................. Periphyton ........................................... Diurnal oxygen curves ................................ units CC...0.0.0.0000...0.00000000000000000.0.0.000... RESULTS AND DISCUSSION Physical and Chemical Environmental Factors ............... Methyl orange alkalinity and pH ...................... Turbidity and seston 0.0.0.0...OOOOOOOOOOOOOOOOOOIO... Turbidity and absorbency ............................. Primary Production and Energetics ......................... Periphyton oooooooooooooooooosooosooooooooooooooooooooooooo. Optical characteristics of pigment extracts ........... Phytopigment - weight relationship ................... Periphyton production ................................ Diurnal gas curves ................................... Energy conversion .................................... Nutrient Supply and Utilization ........................... Phosphorus .COOCOOOOOOOOOOOOOOOOO0.000000000000000000000000. Total phosphorus ...................................... Soluble phosphorus .................................... Biotic utilization of phosphorus ...................... Phosphorus as a limiting factor ....................... Nitrogen .OOOOOOOOOO0.0.00......OOOOOOOOOOOOOOOOOOOOOO0.0... mania nitrogen OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOI Nitrite plus nitrate 0.0.0.0...OOOOOCOOOOOOOOOOOOOOOOO. Biotic utilization of inorganic nitrOgen .............. iii PAGE «3 0\ tr +4 WNGQNNNN rare 15 16 16 21 30 31 31 37 h6 53 Sb 5? a 72 73 7h 80 TABLE OF CONTENTS (Continued) POllution .OOCOOCOCOOOOOOOOOOOIOO0.00.00.00.00...OOOOOCOOOOOO. C/W and P/W quotients sm'}iA-RY 0.0.0.0000...00.....0..0.00000000000000000000000...... LITERATURE CITED iv PAGE 85 86 91 9S TABLE 1. 2. 3. h. 5. 6. 7. 8. 9. 10. ll. 13. 1h. 15. 16. LIST OF TABLES The turbidity, pH, and methyl orange alkalinity of water samples COlleCted at Dabie Road ooooosooooooooooooosoooo The seston content of water samples collected at Dobie Road during the July 1957 flood ........................ The absorbancy of water samples collected from Dobie Road at various spectral ranges ........................ Net primary production determined by periphyton accrual on artificial substrata ................................ A comparison of observed net production rates and net production rates calculated from the phytopigment - weight regreSSion 0.00.0...OOOOOOOOIOOOOOOOOOOOOO0.0000. The conversion efficiency of surface solar energy into periphyton crop ........................................ Computed.month1y phosphorus loads for the Dobie Road study flea .OOOOOOOOOCCOOOOOOOOOOOOOOOOOOOIO0.0.0...... Peak phosphorus loads for the Dobie Road study area dur- illg fIOOd periods OCOOOOOOOOOOOOOOOOOO0......0.0.0.0.... A comparison of phosphorus uptake by periphyton and net periphyton production .................................. Seasonal phosphorus efficiency quotients calculated for 1% meters or river 0......O...0....OOOOOOOOOOOOOOOOOOOO Seasonal phosphorus/weight quotients of periphyton col- 1ected from artificial substrata ....................... Computed monthly inorganic nitrogen loads for the Dobie Road Study area 0.....OOOOOOOOO0......OOOOOOOOOOOOOOOOOO Nitrogen/weight quotients of periphyton collected from artifiCial substrata ...........0......COCOCCOOOCOCCOOCC Nitrogen efficiency quotients calculated for 100 meters Of river 00.0.0000...OOIOOIOOOOCOOOOOOOOOOOOOOOOOOOOOOCO Nitrogen/phosphorus quotients of periphyton collected frm ”tifiCial substrata .......................C...... A comparison of c/w quotients obtained from a clean-water zone, a recovery zone, and a polluted zone .............. PAGE 19 22 26 39 h8 58 59 61 67 69 78 81 82 83 89 FIGURE 1. 2. 3. h. 5. 6. 7. 8. 9. 10. ll. 12. LIST OF FIGURES Plexiglass substrata and supporting devices ............ The relationship between discharge and.methy1 orange a1- kaliriity fOIIOWj-ng two fIOOd periOds .OOOOOOOOOOIOOOOOOO Changes in turbidity, seston, and discharge during the Jllly19s7f100d........................................ The relationship between suspended.matter and the ab- sorption properties of unfiltered river water to three SpeCtral ranges COOOOOOOOOIOOOOOIOOOOOOOOOOOOOOOO0...... A correction graph for converting experimental data to absorbency values (phytopigment units) related to pig- ment concentration ...............OOOOCCCOOCOOOOOOOCCCOC The relationship between phytopigment units and the ash- free dry weight of periphyton .......................... Seasonal periodicity of net periphyton production in the Red cedar River 0.00.00.00.00...OOOCOOOOOOOOOOOOOOOOO... Primary energetics in the Red Cedar River .............. Phosphorus concentrations of water samples collected at Debie Road .OOCOOOOOOOIOOOOOOOOOOIOOIOOOOOOOOOOOOOOOOOOO The relationship between phosphorus fixation by periphy- ton and periphyton growth .............................. Seasonal variations in the phosphorus/weight quotients obtained from artificial substrata ..................... Nitrogen concentrations of water samples collected.at DObie RoadCCOOOICOOOO0..0.0.0...OCOOOOOOOIOOOOOOO0...... PAGE 10 18 2h 28 33 36 51 56 63 71 77 IN TRODU CTION In the study of community dynamics it is necessary to differentiate between the transfer of energy and the transfer or circulation of materi- al. According to the Second Law of Thermodynamics spontaneous energy transfer (exothermic reactions) is always characterized by an enthalpy change accompanied by an increase in entropy. Qualitatively, an enthalpy change may be defined as the energy absorbed during a transfer process, and entropy as the ungailable energy in a thermodynamic system. There- fore, as the entropy of a system increases its capacity for spontaneous transfer is diminished. The number of spontaneous transfers possible within a system depends upon the entropy increase incurred at each suc- cessive transfer. When applying this law of thermodynamics to energy- flow models based on trophic structure, enthalpy change corresponds to the amount of energy transfered to trophic level n from trophic level n-l. Similarly, the entropy change corresponds to the energy changed to heat by the respiration of trophic level n. In constrast to energy-flow, basic materials such as carbon, phos- phorus, and nitrogen.may undergo an infinite number of transformations and are freely recirculated among all trophic levels until lost by export or entombed by sedimentation. Even under such circumstances it is pos- sible for the same material to re-enter the bio-dynamic cycle by import or changes in the redox potential of the postdepositional environment. Flowing waters appear to be the simplest expression of material circula- tion. Recirculation and sedimentation are kept to a.minimum.by downstream transport. On an annual basis only material retained by organisms having a life cycle greater than a year is available for recirculation. There- fore, in the long run import is approximately equal to export and recir— culation of material among various traphic levels is temporary and minimal. Periphyton is virtually the only autotrophic community in the Red Cedar River. It is also the only aggregation of organisms that can di- rectly extract large quantities of dissolved material from the water. Much of the stream drift that becomes available for filter feeders origi- nates from this community. In other words, periphyton serves as the pri- mary vehicle for the transfer of energy and material. Therefore, the maximum biological productivity of the environment is ultimately deter- mined by the efficiency of the periphyton community in utilizing avail- able solar radiation and dissolved nutrients. This investigation is an attempt to describe energy—flow and materi- al circulation at the primary level in a warm-water stream. The findings are first presented as specific data and then expanded to fall within the framework of the principles stated in the introduction. DESCRIPTION OF STUDY AREA The Red Cedar River is a wam-water stream located in the south cen- tral portion of the Lower Pennisula of Michigan. The stream originates as an outflow from Cedar Lake ('1‘. l N; R. h E; Sec. 28, 29; Livingston Co.) and flows for 119 miles before its confluence with the Grand River in 1 the city of Lansing. The river has a gradient of 2.5 ft mile- and a drainage area of 1475 miles 2. The climatological and edaphic features of the watershed have been summarized by Meehan (1958). Brehmer (1958) made a detailed study of the physical, chemical, and biological characteristics of 5.3 miles of stream located 111 miles upstream from the confluence. Most of this study was conducted at Dobie Road which is located 10 miles upstream from the mouth of the river. At base flow this area has a mean width of 72 ft, a mean depth of 1.6 ft, a mean cross sectional area of 110 ft2 , and a discharge of about 65 ft3 sec-1. The stream bottom is composed predominately of sand with occassional patches of gravel. A thermograph was maintained by the Department of Fisheries and Wildlife, hichigan State University at this station. A detailed summary of the water temperature: present during this study is reported by Brehmer (op. cit.). Pollution studies were made at stations located 25 and 50 meters downstream from the outfall of the Williamston sewage treatment plant. METHODS AND EQUIHViENT pH and alkalinity. pH was measured with a Beckman Model N portable pH meter. Alkalinity determinations were made in the laboratory using the method described by Welch (19L8). Turbidi y. Turbidity was measured on a Klett-Summerson photoelec- tric colorimeter which had been calibrated with a Jackson Turbidimeter. The Fuller's earth standards used in the calibration were prepared in accordance with instructions given in "Standardlflethods for the Exami- nation of Water, Sewage, and Industrial wastes" (APHA, ANNA, FSIWA 1955)- Readings were taken using a.number h2 filter (blue) having a spectral transmission of hOO - L65 mu. Corrections for intrinsic color were made by adjusting the colorimeter scale to zero with a water sample that was filtered through.a.Nillipore membrane (HA type, o.L5 u pore size). §g§£g§. Crude estimates of seston were made by filtering 500 ml of river water through a Gooch crucible equipped with an asbestos filter mat. The crucible was then dried, weighed, ignited, and reweighed; thereby measuring both total and organic seston. Absorbancy. The absorption properties of unfiltered river water to three spectral regions were studied in the laboratory using a Klett- Summerson colorimeter. The filters used were; numbers<1 h2 (blue), 5h (green), and 66 (red) having the approximate transmission.ranges of LOO - h65, 500 - 570, and 6h0 ~ 700 mu respectively. Phosphorus. Total phosphorus and total dissolved phosphorus were measured using methods modified rrdm Ellis, Westfall, and Ellis (l9b8). The procedure differed in that the digested sample was split in half and the pH adjusted to the phenolphthalein end point with saturated NaOH before the color producing reagents were added (Taylor 1937). The <1 Klett-Summerson Photoelectric Colorimeter - Industriallflanual IClett Manufacturing Co., N.Y. absorbancy of the colored solution was measured on a Klett-Summerson colorimeter equipped with a number 66 filter (red). Prior to digestion the water samples used for the total dissolved phosphorus analyses were filtered through an HA type Millipore membrane having a pore size of O.h5 u. Nitrogen. Ammonia nitrogen was determined using the distillation method given in "Standardlfiethods for the Examination of water, Sewage, and Industrial wastes" (APHA, AWWA, FSIWA 1955). Oxidized forms of nitrogen (N02 + N03) were measured using the reduction.method described in the same source. Color produced by neselerization in both of the above techniques was measured with a Klett-Summereon calorimeter using a number 5h filter (green). A micro-Kjeldahl procedure similar to the one described by Belcher and Godbert (19h5) was used to determine the organic nitrogen content of periphyton. The catalyst mixture differed from the one given by these authors in that CuSOh was substituted for HgSOh. Similarly, N325 was eliminated from the procedure. These changes permitted the nesslerization of samples having minimal nitrOgen concentrations. Periphyton. Periphyton productivity was studied by suhnerging artificial substrata in the stream. The substrata were plexiglass 2 when plates, 7 mm thick, having an exposed surface area of 1.h dem fastened to a horizontal crossbar. The crossbar was bolted to a steel post in the stream or attached to a vertical upright which was wedged into a concrete block (figure 1). Experiments conducted by the author and Dr. Morris L. Brehmer showed that periphyton samples collected from plexiglass plates exhibited considerably less variation than samples taken from wood shingles, glass microscope slides, and cinder bricks. .moofi>ee merAOQQSm can mpmppmnsm mmeawfixead .H oaswwm 11 The submersion period ranged from one to three weeks depending upon the accrual rate. The substrata were transfered from the stream to the laboratory by means of individual plastic bags and then frozen. The freezing aided in releasing the growth frem the plastic and facilitated pigment extraction by rupturing the cell walls. The relationship between the absorbency of extracted phytopigments and periphyton weight was studied by using the following technique. First, the growth was removed from the substrate by scraping and then washing it with 95% ethanol. Any macro-fauna, such as aquatic insects, were removed from the solution. The particulate fraction of the sample was separated from the extracted phytopigments by filtration through a Gooch crucible. The contents of the crucible were washed with dilute HCl followed by dis- tilled water. The crucible was then dried to constant weight and organic weight estimated by loss on ignition. The absorbancy of the ethanol sol- uble pigments was determined with a Klett-Summerson calorimeter equipped with a red filter (transmission 6&0 - 700 mu) after the solution volume was adjusted to 50 ml. The solvent was evaporated from the extract and the weight of the organic residue determined by loss on ignition. The weight of the residue was added to the organic weight of the particulate fraction. This sum is the weight estimate obtained from one substrata. In the preceeding technique constant weight is defined as two con- secutive readings of‘: 0.5 mg after an interval of 12 hours. Productivity studies involving the use of artificial subtrata are not new. Cooke (1956) gives a comprehensive literature reviewion the subject. However, experiments (Grzenda 1955, Alexander 1956) directed by Drs. R.C. Ball and Frank F. Hooper were the first to cambine phytopigment 12 techniques used by Kreps and Verjbinskaya (1930), Harvey (l93h),luanning and Juday (19h1), and others with artificial substrata (cedar shingles, and cinder blocks) methods. Diurnal oxygen curves. Primary production was measured using the diurnal oxygen curve method as given by Odum (1956). The rate of change in dissolved oxygen for an area where there is no drainage accrual is dependent upon three component rates; production, diffusion, and respira- tion. Before a gross production estimate can be made from a gas curve it is necessary to estimate both community respiration and diffusion. Odum (op. cit.) gives several.methods for computing each of these rates. Therefore, the specific procedures used in this study will now be mentioned. Odum (op. cit.) states the rate of gaseous transfer into water can be expressed by the relation. D - K S where D a the diffusion rate per area (g 02 m"2 hr-l). K - the gas transfer coefficient at 0% saturation S - the saturation deficit between water and air. The gas transfer coefficient (K) can be calculated using the following expression. K'Z(qm-qe) S -S m 0 where Sm - the predawn saturation deficit. Se - the evening saturation deficit. 13 z = the mean depth (m). q = the rate of oxygen change in the morning (g 02 III-3 hr-1)e q 3 the rate of oxygen change in the evening Using the above relationships a correction for either inward or outward diffusion can be applied to each point on the gas curve. Respiration was indirectly measured during darkness by subtracting the diffusion rate from the observed rate of change to obtain the estimate 3 -1 in g 05 ml hr . This method, like all respiration estimates made in the darkness,assumes that community respiration is constant throughout the day and night. In this study the gas curves were constructed using the "spot dye" method where the data carries the dimensions of g 02 m-3 hr . Gross -2 -l m day is obtained.by'multiplying the mean - —l 111 3day )0 primary production in g O2 depth (m) by the area under the diurnal gas curve (g O2 Odum.(19S6) also gives equations suitable for calculating gross primary production when the rate is expressed as a difference between stations rather than a change per hour. All dissolved oxygen determinations were made using the unmodified Winkler procedure as stated in "Standardlfiethods for the Examination of water, Sewage, and Industrial Wastes" (APHA, AWWA, FSIWA 1955). Units. The dimensions on all measurements presented in this thesis are in the exponential form. Using this system cubic centimeters of 02 per 3 2 square meter per day would be expressed as cm 02 m" day-1. In the lit- erature, particularly the European, chemical concentrations are very often expressed in terms of free radicals (i.e. 302, N03, etc.). In order to make comparisons among various authors all data is converted so that concentrations are expressed in terms of the essential element (i.e. mg -l N.NO 1 . 3 RESULTS AND DISCUSSION Physical and Chemical Environmental Factors 15 16 M.3. alkalinity and pH. The methyl orange alkalinity of water samples collected from the Red Cedar River is shown in table 1. Phenolphthalein alkalinity was observed only once, indicating carbonate is present pre- dominately in the half-bound form (X(HC03)2). At base flow m.o. alkalin- ity was in the vicinity of 250 ppm; however, during the 1957 flood a low" of 112 ppm was recorded. When large fluctuations in discharge occur in a short time span, a plot of m.o. alkalinity vs. discharge approximates an inverse linear or curvilinear regression (figure 2). If a similar plot is made for the year this relationship is obscured by the scatter of the points. This variation is caused by seasonal changes in the absolute bi- carbonate content of the river. The pH values for the year ranged between 7.5 and 8.3 (table 1). No difference was noted among determinations obtained during periods of high and low productivity, demonstrating the photosynthetic utilization of hi- carbonate does not contribute to large pH changes in the Red Cedar. This is to be expected because a high bicarbonate content functions as a pH buffer. The greatest fluctuations in pH were observed during floods when the values approached neutrality. It was shown in the preceeding para- graph that high discharge causes a dilution in the bicarbonate concentra- tion. In water like that of the Red Cedar, containing only bicarbonate, carbonic acid, and free 002,the pH will be determined largely by the OE- arising from the hydrolysis of H00; (Ruttner-19S3). Therefore, the drop can be directly attributed to the dilution of the OH- concentration. Turbidity and seston. A summary of turbidity measurements, taken at Dobie Road, together with discharge data is shown in table 14 Maaimal turbidities were recorded during times of high discharge. However, con- tinous periods of maximal discharge were not accompanied by sustained high Figure 2 e The relationship between discharge and methyl orange alkalinity following two flood periods. 17 - «p -,-—n.- .-_.- Methyl Orange Alkalinity (ppm) 280 260 2140 220 200 180 l60 11:0 120 18 boo 800 | l I La 1200 1600 Discharge (ft3 sec‘l) L 2000 2b00 TABLE 1 THE TUEBIDITY, pH, AND METHYL ORANGE ALKALINITY OF WATER SAMPLES COLLECTED AT DOBIE ROAD Collection M.0. Alkalinity Turbidity Discharge <1 Date pH 4(ppm) Units (ftBSec-l) 1957 3 July __ _. 2‘u 92 ll Juky ___ ___ 200 706 12 July 7.5 112 238 1,9h0 13 July 7.6 138 180 2,290 lb July 7.? 166 7b 1,7h0 15 July 7.8 182 85 1,290 16 July 7.9 20h 65 1,020 17 July 8.0 221 73 728 19 July 8.0 250 73 hoo 22 July 8.1 265 73 259 25 July 8.0 279 82 218 29 July 8.1 279 hl 139 31 July 8.0 290 28 111 6 Aug 8.1 268 22 68 9 Aug 8.2 280 19 66 16 Aug 8.3 269 13 5h 27 Aug 8.1 272 lb 50 3 Sept 7.9 250 9 66 10 Sept 8.0 250 7 h8 16 Sept 8.1 270 7 5h 23 Sept 7.9 270 7 h6 1 Oct 8.1 26h 6 hl 1h Oct 8.0 282 15 5h 25 Oct 7.9 220 22 228 16 Nov 8.0 298 39 h73 27 Nov 8.0 2h0 15 1h2 5 Dec 7.9 278 25 %* 116 11 Dec 8.1 250 17 * h2 17 Dec 7.8 270 13 * 130 19 Dec ___. .___ 177 262 (continued next page) TABLE 1 (Continued) Collection H m.o. Alkalinity :Turbidity Discharge 21 Date p (ppm) Units (rt3sec'1) 1958 7 Jan 7.9 202 65 * 18h 1h Jan 7.9 270 ll * 128 21 Jan 8.0 2hh 7 * 163 28 Jan 8.0 216 15 * 111 h Feb 8.0 220 17 * 92 6 Feb 8.0 280 13 * 100 18 Feb 7.8 272 15 * 68 21. Feb 7.7 261. 22 * 81 27 Feb 7.8 211; 51 ** 111 1 Mar 7.9 131: bl *** 7M: 3 mar 8.0 172 23 5&0 6 Mar 8.0 200 25 1:00 13 Mar 8.0 226 9 2175 19 mar 8.1 210 12 209 25 Mar 8.3 236 16 190 30 mar 8.2 230 1b 162 3 Apr 8.1 2h6 15 lbs 8 Apr 8.0 21414 22 298 11 Apr 8.1 238 12 198 13 Apr 8.1 2h0 16 238 21 Apr 8.2 236 20 lh8 30 Apr 8.2 21.2 20 128 7 may 8.2 262 21 111 12 may 7.9 278 lb 86 26 may 8.1 266 ' 10 h2 2 June 8.0 22b 19 59 5 July 7.? 150 175 560 7 July 7.8 1911 78 391 Q discharge data courtesy of the USGS. * ice cover. H bank ice only. 7‘88? ice jam in study area. turbidity. In other words, high turbidity is dependent upon both the discharge at the ttme of measurement and the duration of maximal dis- charge prior to sanpling. Turbidities recorded at the onset of a flood are significantly higher than those taken at the latter stages, even though the discharge is approximately the same. This effect is well demon- strated by the data shown in figure 3. This suggests the maximal tur- bities that intially accompany floods result from the erosion of stream deposits. The sudden drop in turbidity and seston that occurs in spite of continued high discharge can be interperted as an exhaustion of mate- rial from this source. The seston content of water samples collected during the July 1957 flood is shown in table 2. As expected, high seston concentrations were accompanied by high turbidities. However, variations in the limited data were too great to eastablish a quantitative relationship between the two measurements. Judging from seston and alkalinity measurements, total solids in the Red Cedar vary between approximately 200 and 300 ppm. iMcNamee (1930) re- ports the regional differences in the total solids content ofWMichigan waters. The range in 29 Lower Pennisula river systems was 200 to 500 ppm. Similar measurements for 13 river systems in the Upper Pennisula varied between 100 and 200 ppm. Turbidity and absorbancy. The relationship between turbidity and the absorption of various light fractions is shown in figure h. These curves demonstrate that suspended.materials act as a selective filter favoring the transmission of long-waved radiation. Within the turbidity range of 0 - 20, the absorbancy values for red light (6140 - 700 mu) change little. However, after a turbidity of of approximately 25 units is reached, the TABLE 2 THE SESTON CONTENT OF WATER SAMPLES COLLECTED AT DOBIE ROAD DURING THE 1957 FLOOD Collection Wtal Se'ston Organic Seston Date (mg 1'1) (he 1'1) 11 July 95 18 12 July 111; 18 13 July 99 12 lb July 114 6 15 July 23 7 16 July 5 2 17 July 12 2 19 July 15 3 22 July 10 § 25 July 10 Figure 3. Changes in turbidity, seston, and discharge during the July 1957 flood. 23 -2 u-——- _n__. ‘ Hn—nva— v- - «II-— .1. o 250 200 - 150 .. 100.4 2h Turbidity Units Discharge (ftBsecd ) _‘fl 100—. 80— 60—— hOJ 20_ ma 0—0 Total Seston (mg 1-1) -1 H Organic Seston (mg l ) 2‘5 rel absorption curve breaks into a gentle slope that is roughly linear. In contrast, very slight changes in suspended matter, even at minimal turbidities caused an immediate increase in the absorption of blue light (1400 - 160 mu). The absorption curve for green light (500 a 570 mu) is intermediate in position and behavior to red and blue light. Pietenpol (1918), in a laboratory study of lake water, concluded that suspended matter acts as a non-selective light filter. Similarly, Welch (1952) states that suspended materials in certain turbid rivers non- selectively screen all visible wavelenghts of light. Birge and Juday (1930) showed that heavily stained or very turbid lake water favored the transmission of long-waved radiation. They also reported that increasing transparency resulted in the selective transmission of increasingly shorter waved radiations. Likewise, Ellis (1936) found that very turbid rivers were slightly selective in the transmission of longawaved radiation. In the Red Cedar River, even at maximal transparency, red transmission always exceeded green and blue, and green transmission was always intermediate between red and blue. In interperting the data shown in figure h it must be remembered that the system contains two factors that contribute to light extinction. They are; the suspended particles, and dissolved color. water is not considered a factor because the calorimeter was adjusted so that trans- mission was 100% when it contained a distilled water blank. The absorb~ ancy values for filtered water samples (HA typelflillipore membrane, O.hS u pore size) ranged from nearly zero to about 0.06. However, variations functioned independently of turbidity. Therefore, suspended matter is the controlling extinction factor and dissolved color is an accessory factor that introduces random errors into the experimental data when TABLE .3 THE ABSORBANCY OF WATER SAMPLES COLLECTED FROM DOBIE ROAD AT VARIOUS SPECTRAL RANGES Collection Absorbancy Date Spectral Range boo-1.65 mu 500-570 mu 610-700 mu 1957 3 July 0.111 0.03h 0.01b 11 July 0.586 0.356 0.228 12 July 0.777 0.h9h 0.3h6 13 July 0.huh 0.260 0.150 18 Ju1y 0.292 0.126 0.066 15 July 0.338 0.150 0.092 16 July 0.268 0.102 0.086 17 July 0.276 0.108 0.050 19 July 0.276 0.108 0.0h6 22 July’ 0.276 0.130 0.060 25 July 0.202 0.070 0.036 29 July 0.15h 0.060 0.038 31 July 0.11h 0.086 0.02h 6 Aug 0.092 0.036 0.012 9 Aug 0.080 0.036 0.018 16 Aug 0.066 0.022 0.010 27 Aug 0.060 0.020 0.006 3 Sept 0.056 0.022 0.010 10 Sept 0.038 0.018 0.010 16 Sept 0.03h 0.020 0.010 23 Sept 0.080 0.018 0.008 1 Oct 0.08h 0.018 0.006 1h Oct 0.02h 0.012 0.006 25 Oct 0.112 0.058 0.028 16 NOV 0.23h 0.096 0.0h8 27 Nov 0.106 0.03h 0.012 5 Dec 0.098 0.036 0.01h 11 Dec 0.110 0.026 0.010 17 Dec 0.090 0.032 0.008 Figure L. The relationship between suspended matter and the absorption properties of unfiltered river water to three spectral ranges Dots represent mean absorbancy values. - 27 Absorbancy 28 0.- 1. 0.7j Spectral Range 0'61 O 6110 - 700 mu + A 500 - 570 mu X 1400 - 1465 mu 0.5— + 0.1;— ‘ l 0.} + 1. O 0.2—~ O 0.1« . O 0 I I I j I I D I I 25 SO 75 100 125 150 175 200 225' 2‘50 Turbidity Units they are presented as an absorbancy vs. turbidity plot. James and Birge (1938) conducted laboratory experiments where the absorption prop- erties of each component of a three factor extinction system (lake water, dissolved color, and suspended particles) were analyzed. Primary Production and Energy Transfer 30 31 PERIPHY TON Optical characteristics of pigment extracts. Experiments showed that the absorption of broad spectrum light (600-700 mu) is not linerly related to the concentration of 95% enthanol extracts of phytopigment. The deviation from the Lambert-Beer Law becomes apparent at an absorbancy of 0.20 when read on a Klett-Summerson calorimeter and increases propor- tionately with higher concentrations. Nonlinearity was also noted for measurements made with monochromatic light at the peak absorption wave- length. Inorganic solutions (Harrvey Standards) did not exhibit nonlinear- ity with either monochromatic or polychromatic light. Therefore, it is believed this effect as due to interaction between the solvent and solute or to changes among the molecules (e.g. polymerization). The measured absorbancy may be corrected to correspond with the theoretical absorbancy as related to concentration by constructing a correction graph (figure 5). This graph is made by plotting absorbancy against concentration as determined by dilution. This portion of the curve corresponds to the line labled EXPERIMENTAL in figure 5 . The line labled LAMBERT-BEER corresponds to the theoretical absorbancy of the solution. The correction graph is used in the following manner. The measured absorbancy is found on the ordinate and is followed horizontally to inter- cept with the experimentally detemined line 3 this intercept is then read vertically to intercept the extrapolated LAMBERT-BEER line; the absorbancy unit opposite this intercept represents the corrected absorbancy reading. In order to avoid confusion between measured absorbancy and corrected absorbancy, the corrected absorbancy will henceforth be designated as Figure 50 A correction graph for converting experimental data into absorbancy values (phytOpignent units) related to pigment concentration. The emeriments were jointly conducted by Brehmer (1958) and the author. 32 Absorbancy 33 5.0 0.1 J _L 1, l | 1 LJ. is l J ILJ1¢ 10 Relative Concentration 100 3h phytopigment units. Phytopigment-weight relationship. Experiments showed that phyto- pigment units could be used to make quantitative estimates of organic weight (ash-free dry weight) for phytopigment values less than 1.3. For greater values the variations were too large to be useful as a quantitative technique. However, such values are still valuable for comparative studies. It is believed that the increasing variation was due to the physical state of the algal colony. More specifically, samples having the greatest variations were those which supported the most luxuriant growth of peri- phyton. The increased variation is probably a combined effect of the death of some members of the colony and the accumulation of organic detritus on the surface of the substrate. Therefore, for quantitative studies it is important that the exposure period be short enough to avoid this condi- tion. The data presented in figure 6 were collected from July 1957 to July 1958. The exposure periods ranged from one to three weeks depending upon the accrual rate of periphyton. A linear regression wassfitted to the phytopigment-weight relation for phytOpigment values less than 1.3. The model used was Model I as given by Snedecor (1956). The sample regression for this model is A Y - a + bx where in this case the mean weight estimate. the intercept of the Y axis. the point estimator of the population slope. the observed phytopigment reading. xdwK> IIII .35 mmpmfipno Hmzpfifidca 98 A98 nmpmfinpmo some you Eponm m.“ seamen 8:03.300 Rmm one .cofnnaaama mo “Emacs S oonmnnmm one. new and? psoewunaofinna c0258, mEmcowpmHmu one .o shag 36 3.3.: pcmaawwaofmnm m; «A a; 04 a. m. a. o. m. a. A _ a a _ _ A ON 3 00 om OOH QNH l 9: 00H ( 3m) 111819“ 37 Using the computed constants the equation for predicting 4% from X becomes 1% - - 1.72 + 82.37 x Equations for computing a confidence region on the regression are given ' by Snedecor (1956). The confidence region appears as curved borders on both sides of the regression. The width of the confidence belt for any prediction made by X is dependent upon how far removed X is from i (the mean of the X's). Since all of the values of X in figure 6 are close to i (0.52), the confidence region appears as. two lines that are nearly parallel. The mandmum 95% confidence limit for any estimate of Y (indi- vidual estimates) made from phytopigment values between 0 and 1.3 is i 29.03 mg and is located at the mandmum value of X. The minimum 95% limit is found at i, and in this case it is I. 28.h2 mg. The same limits for I (mean estimates) are _4_-_ 3.05 mg and i 6.82 mg respectively. Periphyton Lroduction. Net periphyton production was measured, using artificial substrata, from July 1957 to July 1958. The taxonanic composition of the camunity varied with the season. Gomphonena, Navicula, Diatoma, and Fragellaria were the most common genera in spring and summer communities. Cocconeis was the dominant genus in the early fall. Late fall and winter communities were composed almost entirely of Navicula and Fragellaria. Peters (1958) gave a very detailed account of seasonal diatom periodicity in the Red Cedar River. Peters also found that the taxa present on the stream bottom and natural substrata were the same as those colonizing artificial substrata. Similar observations were reported by Butcher (1932) and Patrick, Horn, and Wallace (1991) for periphyton colonizing glass slides. The seasonal peridicity of net periphyton production is shown in 38 figure7'. The mean rates ranged between 0.11 and 22.78 mg ash-free dry weight dew-2 day-1. The highest rates were observed in June and the lowest when the river was covered with ice. In the latter part of August production dropped very sharply. In lessathan a week the mean rate fell from 9.51. to 0.52 mg dcm'2 day'l. This situation was observed at the same time by Brehmer (1958) for sim.stations upstream from Dobie Road. There was no great change in the physical (turbidity, temperature, and discharge) or chemical (pH, alkalinity, nitrogen, and phosphorus) charac- ter of the stream concurrent with production cessation. As fall pro- gressed there was a gradual increase in production that subsided with the event of ice cover. Prior to the decline, the community was composed of many genera of diatoms. By mid-September the colony was comprised almost entirely of Cocconeis. This suggests the decline was caused by a shift from a strong light - warm water community to a weak light - cold water community (Ruttner 1953). However, the abruptness of produc- tion cessation is still very puzzling. E.P. Odum (1959) has adjusted and corrected primary production estimates obtained by various methods and different authors to make than comparable with one another. Using these conversions he gives annual net production estimates for Silver Springs, Florida (H.T. Odum 1957) and the Sargasso Sea (Riley 1957). They are 7.h and 0.26 g dry organic weight m-2 day.1 respectively. Data given by H.T. Odum (op. cit.) and Riley (op. cit.) suggest that in aquatic biotypes gross produc- tion is approximately equal to two times net production. On this basis the mean annual gross production rate in the Red Cedar River would be about 1 g m-2 day-1. This places the Red Cedar River in the second order of productivity along with grasslands, coastal seas, some shallow 39 TABLE 0 NET PRIEARY PRODUCTION DETERMINED BY PERIPHYTON ACCRUAL 0N ARTIFICIAL SUBSTRATA Collection Exposure Water Temp. {Jean Two Standard Date Period Range for Net Production <3. Errors (days) Period 00 (mg dcm'2day’1) (rg dcm'zday‘ 1957 (i) 31 July 7 18.0-22.5 3.28 1.13 8 Aug 7 18.0-22.0 6.58 0.8h 16 Aug 7 18.0-2h.0 9.5L 1.53 27 Aug 7 1705.200; 0.52 0.16 3 Sept 1h 18.5-2200 2006 0017 17 Sept 1h 1h.0-18.0 1.0a 0.12 1 Oct 1L 1h.0-18.0 1.25 0.20 18 Oct 1b 6.0-1h.5 2.10 0.89 b Nov lb 6.0- 8.0 b.08 0.32 5 Dec 1L 0.;- 1.0 0.86 0.2h 1958 7 Jan 3h 0- 6.5 0.13 0.08 28 Jan 21 0 0.11 0.02 18 Feb 21 0 0.12 0.05 19 Mar 28 0- 6.5 1.01 0.26 17 Apr 114 700.1700 8.33 0071 12'Nay’ 7 10.0-17.0 11.L7 1.22 19 May 7 11400-2100 5011 00 S7 11 June 9 1b.0-20.0 11.3h 0.71 18 June 7 11100-2000 21021 70135 25 June 7 1h.0-20.0 22.78 1h.72 To convert the data to g m"2day"1 move the decimal point one place to the left. b0 0 OH O p H “fl flu é 93mg bl hash ecsw be; ha< aw; new new ova >02 #00 pmom ws< has: 4H mm as am ea mm as mm ea mm ea mm 4H m. as . c mm ea mm 4H mm H z _ _ up, _ _ A: .r u_ _ ea. _ __ _ z m _ww dd mm AH __ ifi OH NH 4H 0H ma ON NN em _AEp a_mop 8m) notionpoad uoafludtaed 19M I ( L2 lakes, and ordinary agriculture (E.P. Odum 1959). A comparison of net production rates determined by loss on ignition and phytopigment methods is shown in tabley5. The degree of agreement between the two methods is remarkable considering the regression was computed from diatom populations having different taxa living under various light, temperature, and nutrient conditions. These findings are inconsistant with some previous investigations. Ryther (1956) points out that many workers have shown that chlorophyll content per cell varies over a wide range depending upon the physical and chemical environment. However, Tucker (19h9) demonstrated a good correlation (0.8h) between planktonic diatom counts and pigment absorbancy} Similar findings were reported by Peters (1959) for diatoms in the Red Cedar River. The use of artificial substrata, like other cr0p methods of measuring net production, has intrinsic shortcomings which tend to lower the rate estimate. They are: predation by consumer organisms; and in the case of stream periphyton, downstream drift. Both of these sources of error can be held to a minimum by removing the substrate when there is a firm uniform coat of periphyton. Longer exposure results in a flocculent colony that is susceptible to losses by downstream drift and insect pre- dation. As a general rule, this condition can be avoided by removing the substrate before the colony has an ash-free dry weight of 150 mg. It is not likely that drift and predation were significant in this study except in three instances when production was maximal. At this time the substrata were clearly over exposed since they supported a flocculent colony densely populated with aquatic insects. At other times the sub- strata supported few if any insects and required vigorous scrubbing to remove the algal coating. TABLE 5 A COMPARISON OF OBSERVED NET PRODUCTION RATES AND NET PRODUCTION RATES CALCULATED FROM THE PHYTOPIQiENT-WEIGHT HESRESSION Collection Observed Mean Rate Calculated Mean Rate <1 Date (mg dcmfzday'l) (mg dcmfzday’l) 1957 8 Aug 6.58 6.81 3 Sept 2.06 2.53 1? Sept 1.0h 2.22 1 Oct 1.25 1.65 18 Get 201).; 1.77 h Nov b.08 3.28 5 Dec 0.86 0.30 1958 7 Jan. 0.13 0.30 28 Jan 0.11 0.02 18 Feb 0.12 0.05 19 Mar 1.01 1.32 17 Apr 8033 7009 5 May 10.12 11.20 12 May 11.h7 15.5h 19 May 5.11 6.21 29 may b.89 5.75 18 June 21.21 19.7h 25 June 22.78 16.38 <1 Calculated rates computed from Y I ~l.72 + 82.37 X where Y - mean weight estimate X - mean phytopigment reading hb Diurnal gas curves. Sargent and Austin (19h9) used changes in dissolved oxygen to estimate the productivity of a coral reef. Their methodology did not correct for oxygen gains or losses due to diffusion. However, this was not critical because deviations from saturation were small relative to total oxygen production. Odum (1956) gave a.more sophisticated version of this technique that provided for diffusion cor- rections. This method was successfully used by Odum (op. cit., 1957a) in Florida springs and a turtle-grassscommunity; and by Kohn and Helfrich (1957), for a coral reef. However, Mchnnell and Sigler (1959) found that Odum's method was unsuitable for productivity studies in a shallow rapid river. Diurnal oxygen curves were found to have only limited value in the Red Cedar River for measuring primary production. One test trial out of four was successful. Two trials were conducted during the winter when production was minimal; both were unsuccessful because no change in dis- solved oxygen could be detected between the upstream and downstream sta- tions. Iieasurements made during a period of high productivity yielded paradoxical data; that is, negative changes occurred at dusk and dawn that exceeded the calculated losses attributed to diffusion and community respiration. This indicates the methods for computing the gas transfer coefficient (K) and respiration were inadequate. The formation of oxygen bubble by photosynthesis could be respon- sible for-these and other errors inherent to gas curve productivity measurements. Bubble escapement to the surface would result in under- estimates if such losses were not considered by the investigator. Odum (1957a) reckoned with this problem and was able to measure the net pro- duction of benthic algae by collecting ascending bubbles with a funnel b5 trap. Such a technique asswmes that the bubbles are lost to the surface shortly after formation. This was not the case in the Red Cedar River. At times bubble accumulation within the algal mat became so great that walking in the stream gave the water an effervescent appearance. This condition was still present after nightfall and absent by the following morning. Presumably the oxygen went into solution at night when the water was unsaturated. Nocturnal solution of oxygen bubbles would intro- duce several errors into the production estimate. The gas transfer co- efficient (K) is calculated on the basis of nocturnal dissolved oxygen changes assuming such changes are caused solely by diffusion and commu- nity respiration. However, if bubble solution takes place (at night) the extent of outward diffusion will be underestimated. This will result in an error in the computation of K. For the same reason the community res- piration.measurement will also be incorrect. Since no quantitative data are available to illustrate this effect it is impossible to say whether such descrepancies would cause other than minor anomalies in the data. However, it is likely that bubble accumulation during the day could cause a significant underestimate simply because large quantities of oxygen did not go into solution when the productivity measurements were made. In April 1958 a diurnal gas curve, free from anomalies, was obtained from the Red Cedar-River at Dobie Road. The gas transfer coefficient -2 -1 (3.1 g 0 m hr ) is in general agreement with literature values quoted 2 by Odum (1956). The gross and net production rates-were found to be 3.6 and 2.h g 02 m’eday-l respectively. Odum.(1957a) reports gross production rates from 0.7 to 6b g 02m"2 '1 production / community respiration quotient for Dobie Road is 3.0, which day in 11 Florida springs. The gross according to Odum (1956) classifies it as an oligosaprobic community} Energy conversion. The energy content of periphyton samples was estimated using chemical analyses and caloric values obtained from the literature. Birge and Juday (1922) report the following analyses for diatoms collected from Lakelfiendota: crude protein, 37.81%; fats (ether extract), 22.h8%; carbohydrates (crude fiber plus nitrogen free extract), 39.71%. These percentages are in terms of the ash-free dry weight. Standard caloric figures on a dry weight basis are; 5,650 calories<1 per gram protein, 9,b50 calories per gram fat, and h,100 calories per gram carbohydrate (Juday 19h0). Solar radiation data were obtained from a pyrheliometer station.main- tained by the Soil and Water Conservation Research Division of the U.S. Department of Agriculture. The measurements express the amount of solar energy received by a horizontal plane at the earth's surface and carry the dimensions of g cal cm'z. The pyrheliometer station is located ap- proximately two miles west of the Dobie Road study area. Lindeman (l9h2) defines energy intake efficiency as the ratio of energy intake at a given trophic level to the energy intake at the pre- ceding (It/It-1)' According to Odum (1959) at the primary level this can be expressed as the ratio of gross production to surface radiation (Pg/L) or as the ratio of gross production to available radiation (Pg/La)' Juday (19h0) in addition to using the efficiency percentage Pg/La employed the following efficiencies based on net production (Pu); Pn/L and Pn/La. Seasonal Pn/L efficiencies obtained from the Red Cedar River are shown in table6 . They ranged from 0.003% to 0.216%, with an annual mean (l The calorie or small calorie is a cgs unit of heat defined as the quantity of heat required to raise the temperature of one gram of water from 3.500 to L.s°c (Handbook of Chemistry and Physics. Chem. Rubber Pub. 00., Cleveland, Ohio). In pythIiometry the same unit is called the gran- calorie (Crab 1950). D7 of 0.072. Juday (19LO) estimates that 1/3 of the total energy fixed by photosynthesis is degraded to heat by plant respiration. 0n the basis of this figure the annual mean Pg/L efficiency for the Red Cedar is about 0.01%. This value is lower than most similar values reported in the literature. According to Odum (1957) Silver Springs, Florida has a Pg/L efficiency of 1.2%. Using respiration figures given in the same work it was calculated that Silver Springs has a Pn/L efficiency of about 0.7%. Juday (op. cit.) reports annual Pn/L and Pg/L values for Lakeldendota, Wisconsin of 0.27% and 0.35% respectively. Cedar Bog Lake,liinnesota has a Pg/I efficiency of 0.1% (Lindman 19h2). Using data given by Lin- deman (19hl, op. cit.) it was estimated that Cedar Bog Lake has a Pn/L value of 0.07%. Efficiencies reported by Riley (19h1) and Clarke (l9h6) for productive marine areas were roughly the same as those cited for Lake Mendota. Clarke (op. cit.) points out that efficiency estimates made on the basis of surface radiation are a function of the photosynthetic efficien- cy of the organism and light transmission within the environment. It should be recalled that the Red Cedar is subject to prolonged ice cover as well as high turbidity caused by periodic floods. Under such condi- tions lower efficiencies (Pg/L and Pn/L) are expected in comparison to less turbid lacustrine and marine environments because more energy is lost to abiotic absorption. The Red Cedar has a productivity comparable to most temperate lakes and shallow marine areas (page 38 ) even though the relative utilization of surface radiation is lesse This indicates the Red Cedar is more efficient (Pg/La and Pn/Ia) in converting available radiation into plant products than other biotypes having the same level of productivity. This idea can be clarified by restating Odum's (1959) TABLE 6 THE CONVERSION EFFICIENCY OF SURFACE SOLAR RADIATION INTO PERIPHYTON CROP Collection Exposure Energy Content Surface Radiation % Date Period of Crop for Period Efficiency (days) (g cal cmfz) (g cal cm'z) (Pa/L) 1957 31 July' 7 1.h h,269.9 0.032 8 Aug 7 207 33h5509 0.078 16 Aug 7 307 3,609.0 0.101 27 Aug 7 0.2 3,2h6.h 0.006 3 Sept lb 1.0 5,109.8 0.019 17 Sept lb 0.9 5,518.6 0.016 1 OCt lb. 1.0 5,071.7 0.020 18 Oct 1h 1.8 3,779.2 0.0h6 h N0v lb 3.h 2,326.7 0.1h5 5 Dec 1h 0.? 1,702.9 0.0h2 1958 7 Jan 3h 0.3 h,302.7 0.006 28 Jan 21 0.1 3,563.3 0000).]. 18 Feb 21 0.1 h,839.2 0.003 17 Apr 1h 6.8 6,h92.9 0.105 5 May 10 6.0 5,170.1 0.115 12 may 7 h.7 h.502.0 0.105 19 may 7 2.2 h,6o9.7 0.0h8 29 May 10 2.9 6,713.7 0.0h3 11 June 9 6.0 h,955.3 0.121 18 June 7 8.6 h,21h.8 0.208 25 June 7 9.h 3,838.7 0.2h5 b9 definition of photosynthetic efficiency based on available radiation. (1) Efficiency = Pg/La or (2) Pg = La x Efficiency (3) La = Pg / Efficiency From equation (2) it can be seen that a given level of productivity can not be concurrent with a reduction in available radiation unless the photosynthetic efficiency increases. Odum (1959), on the basis of data given by Rabinowitch (1951), states the overall efficiency of available light fixation by the plant world is about 1%.. There is some justification for extrapolating this estimate to aquatic communities. Juday (l9h0) reports an annual Pg/La efficiency of 0.91% for Lake Mendota. A rough estimate of the annual mean available energy input (La) for the Red Cedar can be made by substituting this figure (1%) and the annual mean gross production estimate into equation (3). The estimated La is 3b.h g cal cm'z day"1 or 10.3% of the mean surface energy input. A summary of primary energetics based on the data and assumptions stated in the text is shown in figure 8. Two categories of abiotic en- ergy loss are recognized in the energy-flow'model. The first is the loss of surface energy by reflection and energy degradation caused by the absorption of transmitted light by the aqueous medium and suspended sol- ids. This loss is calculated by subtracting the radiation reaching the stream bottom from the surface radiation (L - La)' The second is the degradation of available light that is not used in photosynthesis. This quantity is estimated by subtracting the radiation reaching the stream SO .GOAQHHAQMDh I m G .cowposooud_hnmsdua pod I m m .cowaosooadchhoswud”macaw I m m .mwoa hmaodo capofiba Aspen I m I A m w anamonpnhmopogd ca oouwfiaps soc soppon saunas on» mcanomou zmpoco I m a A w .cowmmasmdsap mdwuso cOAodaomna ssAooE odd dowpooamoh an pmoa hwpoco I A I A OHIE N CA oowmoudxo one ounce amazes no woman span a .soppon smoke» and moanomou amused I A .hmuoco oceansm I A lumposavcoo moaneddolsuhm .uo>«m havoc com ens ca mofipowaoco zhmsfipm .MWOAdem 51 soppom seesaw 65.3 goes: .haa A Amm.amv am.ov a ‘ uH O A/Jyx a Amo.sm as u «A Amo.~muv as a a AmA.Hmmv m a n a Aam.AmmV swam 52 bottom (La - Pg). The expression (L - La) + (La - Pg) may be reduced to (L - Pg) and is termed the total abiotic energy loss. Energy lost by evaporation and energy gained by condensation are not considered in the model because they involve energy that has already been converted to heat. The preceeding computations assume that primary production is lim- ited to periphyton on the stream bottom. This assumption appears to be accurate in that extractions from suspended materials obtained by fil- tration of water samples showed a virtual absence of phytopigments. Since only primary energentics are considered, periphyton respiration is the only source of biotic energy loss. The annual mean rate of net production (0.23 g cal cm-2day'1) is the amount of energy that is avail- able for transfer to consumer organisms. The term consumer as used in the framework of this model includes the reducer organisms which are usually considered separately in the more complex energy-flow*models. Nutrient Supply and Utilization 53 Sb PHOS PHORUS The ph03phorus concentration in water samples collected from July 1957 to July 1958 is shown in figure 9. The sampling frequency was dictated by existing stream conditions; that is, periods of change such as floods were sampled daily and static periods were sampled two or three times a week. The concentration data were converted into load estinates. A load is defined as the absolute quantity of detrital and dissolved phosphorus tramported into the study area by suspension in a given period of time. a soluble load refers to tutorials carried in solution. The following terms are used synonymously throughout the text: load with import; and downstream drift with export. Traction and sealtetion transport are not comidered in this study. Investigations on the Red Cedar River by Kevern (ms) itflicxte that phosphorus transport by traction and saltetion is negligible. Krumbein err! Sloss (1956) discuss the classification and characteristics of various modes of stream tramport. Load estimtes were nude in the following way. The observed phos- phorus concentration for the day was multiplied by the total discharge for the same day. This product was then plotted against the sampling date. 5 curve was constructed by repeating this process for all samples taken during the month. The area under the curve was then measured and taken to be the quantity of phosphorus transported into the study area- for that period of time. The accuracy of this method is dependent upon two assumptions: first, that the increase or decrease of phosphorus between the observed points is uniform; second, that all major changes in phosphorus l Master's thesis in preparation, Dept. Fisheries and Wildlife, M.S.U. SS pmom manoa pm popooaaoo mmHaEwm pops: mo m:0fipmupcmocoo mspozmmosm .m Page 56 has“. 023.. he 2 pg. .8 2 new 82. com #02 p00 paom mg. hash. 8 fl 3 fl 2 fl mm fl cm .3 mm i R as mu .2 ma fl 3 fl 3 fl mm fl am i b I L t L #7 _ I P : . _h _ : _ __ . b. b I _ hp _ _r _ r0: 1 1 < 4 C d a . . . .ee eee e @100 . er . ., e e e e .e r e e ee . e Ow . e e e e ee e 18H . r 03 e . of“ kc . fi 03 . f 02 \ m mfiosamoi 85°83 H38. TIL ”Eofimoé H38. O|lo : com com - oom - 8m cam I_ Ham snuoHdsond 3n S7 concentration.were recorded by the investigator. The chief advantage of this method is that an estimate of phosphorus import can be made for any time period simply by measuring the area of the curve over the desired dates. Total phosphorus. The monthly distribution of phosphorus is de- pendent upon the discharge trends of the river. The months of July, December, February, and March accounted for 59.62% of the total discharge and for 65.39% of the phosphorus load for the year; The maximum monthly import was observed in.July'19S7 when 5,992 kg P were transported into the study area. Similarly, a minimum load of 299 kg P as recorded in August 1957. The peak phosphorus loads that accompanied individual floods are shown in table 8 . These data show that h5$ of the annual phosphorus import was carried into the study area by three floods having excombinsd duration of 31 days. Therefore, floods function to make nutrients most available when the conditiom for their utilization are - least favorable. The phosphorus yield of the watershed area is estimated to be 50 kg P mile"2 yearrl. Sawyer (19h?) reports a figure of 72 kg P mile“2 year'1 for agricultural drainage and runoff in the Madison, Wisconsin, lake region. Brehmer (1958) estimated the annual phosphorus output of a sewage disposal plant located 9.9 miles upstream from the study area as 2.5 metric tons. This quantity of phosphorus is 15.5% of the yearly import at Dobie Road. Soluble phosphorus. The soluble phosphorus loads were more uniform than total loads even during periods of high discharge (table 7 ). A mnmum value of 726 kg P was recorded for the month of hrch 1958. Similarly, a minimum value of 162 kg P was observed in November 1957. TABLE 7 CO‘JPUTEH) MONTHLY PHOSPHORUS LOADS FOR THE DOBIE ROAD STUDY AREA DATA ARE GIVEN IN KILOGIAMS OF ELEMENTAL PHOSPHORUS Month Total Percent Soluble Percent Discharge P of Total P of Total (m3 sec‘l) 1957 July oz uoo seem ws< hash. 8 mm 32% a m mam film as as 3.11%.: mm “a _awtfimm a mm in 2.: 7 J o O / l .1 ON .1 . 1 0m W om E01 ‘x (LHDIaM / SflHDHdSOHd) 72 Other times this relation was not so well defined. Perhaps the high quotients (13.02 - 18.27) observed in May and June when production was seminal resulted from over exposing the substrata. It must be remembered that the mean production rates were computed on an arithmetic basis. Therefore, it is quite possible that production at the time of collection was minimal and hence the phosphorus concentration maximl. Such a situation was described by land (l9h9) in his studies of annual spring blooms of Asterionella fomosa. In this case it was observed that the phosphorus concentration within the cell was much greater near or at the peak of a bloom. Nihei (1955) reports a mechanism that explains the increase in phosphorus concentration accompanying growth cessation. Working with Chlorella he found that prior to mitosis there is an accumlation of polyphosphates which are used as an energy source in processes connected with cell division. As division occurs the phosphorus is dissipated- by being redistributed to the new cells. However, if mitosis is inhibited there is an accumulation of polyphosphates within the cell. In January 1958 two sets of substrata: were eXposed for periods of 21 and 311 days respectively. At the time of collection both sets had PM 1: 103 quotients that were practically indentical (33.55 and 33.63). There are two factors which indicate a P/W quotient of this magnitude represents the comunity saturation concentration. It is to be noted that an additional 13 days of exposure failed to cause an increase in phOSphorus concentration; furthermore, production was minimal and hence polyphOSphate accumulation mammal. The periphyton colony was predomi- nantly composed of two genera of diatoms; Navicula, and Fragellaria. Phosphorus as a limityg factor. Soluble phosphorus concentrations 73 found in the Red Cedar River ranged between 21 and 108 ug P 1-1 ; with an annual mean concentration of 50.] ug P 1'1. Chu (19143) working with laboratory cultures gave values for minimal concentrations of phosphorus just permitting optimal growth in six species of phytoplankton. These values ranged from 6 to 29 ug P 1.1. More recent studies, by Rhoda (191.8), indicate that Chu's miniml concentrations are too high when dealing with natural populations. Rhode found that Asterionella fgrmosg1 could thrive in a lake having phosphorus concentrations that were inadequate for growth in artificial media. Fogg and Westlake (1955) suggest that some compound, possibly a peptide, is present in lake water which facilitates the ease of phosphorus utilization. Thus, on the basis of experiments by Lund (0p. cit.), Rhode (op. cit.), and Chu (op. cit., 19115) it seems very unlikely that the phosphorus concentrations found in the Red Cedar are low enough to limit periphyton production. It is more probable that the high level of phosphorus present in the Red Cedar would tend to limit or exclude certain taxes of periphyton. Rhoda (op. cit.) studied the chrysophycids Dinobryon divergens and Uroglena smericana and found they were able to reproduce at a maJdmal rate when the phosphorus concentration of the water was bea-rly detectable. Both Species became inhibited after the phosphorus concentration was raised to 1.6 ug P 1'1. NITROGEN The inorganic nitrogen concentration in water samples collected from 26 November 1957 to 7 July 1958 is shown in figure 12 . The nitrogen sampling schedule, like the one described for phosphorus, was determined determined by the discharge trends of the river. Similarly, nitrogen 7h loads were computed by the area under the curve method. The method of analysis did not provide for the differentiation between nitrate and nitrite. Therefore, the total inorganic nitrogen concentration was accounted for in the following way; nitrate plus ni- trite, and ammonia nitrogen. Nitrite occurs in natural waters as an oxidation-reduction intermediate between ammonia and nitrate and is seldom present in appreciable amounts except in cases of extreme organic pollution. Ammonia nitrogen. Ammonia in natural water is present in the form + of NH,4 and NHhOH. For the usual pH values found in the river the ratio of NHh+ to NHhOH would be approximately 30 to 1. Under normal winter and fall conditions the ammonia concentration never exceeded 0.30 mg N.NH3 1-1. During the summer and spring months ammonia is usually present only as a trace (Brehmer 1958). However, an exception to this general rule was noted following an industrial accident which occurred 35 miles upstream from the experimental area. . 0h.21fiMay 1958 a fire destroyed a portion of a.metal plating plant located in.Fow1erville, Michigan. In the course of extinguishing the fire water flooded the cyanide tanks and toxic chemicals were diverted into a waste lagoon . The lagoon outlet was closed and its contents treated with sodium hypochlorite to oxidize the cyanide to the relatively non-toxic cyanate (Eldrige 1933, Dobson l9h7). Cyanate is hydrolized to ammonia compounds at a rate which is a function of pH. When the reaction was complete the contents of the lagoon were feed into the river. It is thought this incident was responsible for the extraordinarily high ammonia concentrations (between 0.25 and 0.73 mg N.NH3 1'1) observed during the first 1h days of June. 75 The maximum monthly ammonia load was noted in February 1958, when 2,1”; kg N.HH3 were transported into the study area. A minimum monthly value of 51; kg N.NH was recorded for the months of April and May 1958. 3 In June following the industrial accident, nearly half of the total inorganic nitrogen import (2,765 kg N.NH3+N02+N03) consisted of ammonia nitrogen. Ammonia nitrogen comprised 9.83% of the total inorganic nitro- gen load for- the entire study. This value is not a fair estimate of the per annum percentage of ammonia for two reasons. First, the study was weighted in favor of those months characterized by the presence of ammonia (Brenner 1958); second, because of the high concentrations of ammonia at- tributed to the accident at Fowlerville. In any case, it is safe to say that ammonia comprises only a small fraction of the total inorganic nitro- gen that is available for biotic utilization in the Red Cedar River. Nitrite and nitrate. The monthly nitrite plus nitrate loads are given in table 12. These values were computed for a study period ranging fran 26 November 1957 to 7 July 1958. The total import for this period was 75.7 metric tons N.NO +NO3 or 91.2% of the total inorganic nitrogen 2 load. A maximum monthly value of 20.9 metric tons N.I\102+NO3 was noted in December 1957. A minimum monthly value of 1.14 metric tons N.N02-0'NO3 was recorded in May 1958. The nitrite plus nitrate yield of the watershed is estimated to be 314.7 kg N.NO +N0 mile"2 month-1. The total phosphorus yield for the 2 3 same period was h.l kg P mile"2 month-1. The magnitude of these loads, as m the case with phosphorus, was found to be dependent upon the discharge trends of the river. That is, periods of high discharge were accompanied by high nutrient import. However, one essential difference between phosphorus and nitrogen is 76 .pwom 0309 pm popooaaoo moHdEmn pops: mo mcogoppceocoo comoufimz .NH oasmaa 77 haze meow he: un< as: ppm and own >02 3 PM 3 an MA an MA mm ma tom ma on E on m pm mm 4 a o 4 a a o a a o < < E < << < ¢\< o o o e e _ . . w _ a a \ ._ a P M o H a sewage m: .. mmzé l u l mofmoz .z I r_", 1...- (1...! II). m.o o.H o.m TN 49411 ueSoaqtN Sm I- 78 TABLE 12 COMPUTED MONTHLY INORGANIC NITROGEN LOADS FOR THE DOBIE ROAD STUDY AREA DATA ARE GIVEN IN KILOGRAMS OF ELEMENTAL NITROGEN Month NO2+NO3 NH3 Total Inorganic Percent of Nitrogen Total 1957 November <1 1,1L2 132 1,271; 1.52 December 20,850 1,8h8 22,698 27.02 1958 January 13,028 l,bh8 lb,h76 17.23 February 5,316 2,1hh 7,h60 8.88 March 20,592 1,008 21,600 25.71 April 6,098 Sh 6,152 7.32 May' 1,38h 5h 1,h38 1.71 June 1,751 1,01h 2,765 3.29 My <2 5,580 556 6,136 7.30 Totals 75,7h1 8,258 83,999 <1 Data computed for 26-31 November only. <2 Data computed for 1-7 July only. 79 quite clear. A peak load of phoSphorus during a flood is characterized by both a sudden rise and fall. Peak nitrogen loads are characterized by a sudden rise followed by a period of sustaimd nitrogen enrichment. For example, during the February-March flood in 1958 the daily phosphorus loads dropped to their pre-flood levels in four days. However, the daily nitrite plus nitraate loads were sustained at a level eight times greater than the pro-flood values for a period of three and a half weeks. The data indicate that agricultural runoff is the major source of inorganic nitrogen while sanitary drains are the mjor source of phos» phorus. For ample, during an extended freeze in the winter of 1957- 1958 the inorganic nitrogen values drapped uniformly over a: period of a from 2.21 to 0.15 mg N.N02+ NO 3 nitrogen was mintained until a thaw occurred. It is believed this + NH3 1'1. This low level of inorganic decrease my caused by the gradual elimination of agricultural runoff by freezing. During the thaw no depression was noted in the phosphorus concentra- tion. This adds strength to the notion that the origin of phosphorus is sanitary drains rather than agricultural runoff since the former are not subject to freezing. Such an interpéfirtation explains the difference between the nature of the phosphorus and nitrogen loads during and following floods. The rapid and striking rise of both phosphorus and nitrogen would represent the "flushing out" of stream deposits. The rapid recovery of phosphorus to pro-flood levels signifies and exhaustion of phosphorus from this source. On the other hard, the sustained high level of nitrogen would be interperted as: originating from the increased runoff which is maintained for a considerable period after the flood crest has been reached and the stream bed flushed. Prior to the freeze over 90% of the inorganic nitrogen was in the nitrite plus nitrate form with the remainder being ammonia nitrogen. During the freeze when minimal nitrogen concentrations were present nearly half of the total inorganic nitrogen was: ammonia nitrogen. Following the thaw normal winter ratios of ammonia nitrogen to other forms of inorganic nitrogen were restored. Biotic utilizatio_n_ of inorganic nitrogen. The nitrogen content of periphyton colonizing artificial substrata was studied using the previous- ‘Ly cited methods. The N/‘u' x 103 quotients from 5 May to 25 June 1958 are shown in table 13 . The quotients ranged between 9.32 and 22.112. The percentage of nitrogen in the ash-free dry weight of diatomaceous communities in the Red Cedar River varied between 0.937 and 2.211%; with a norm of 1.119%. These values are somewhat lower than those given by Ketchum and Redfiel‘d (19119) for laboratory cultures. They found that the diatom, Nitzschia closteriugg, had a mean nitrogen content of 6.67%. Similar values fer mmbers of the Chlemphyceae averaged between 6.62 and 8.65%. Ketchun and Redfield (op. cit.) also noted that the percentage of nitrogen in Chlorella pyrenoidosa was approximately halved when grown in media deficient in nitrogen, or nitrogen and phosphorus. Algae cultured in phosphorus deficient media had nitrogen concentrations slightly higher than those grown in phosphorus rich media. The net efficiency quotients calculated for 100 meters of stream are shown in table 114 . These values varied from 0.029 to 0.187%. Six out of seven of the net nitrogen quotients were lower than the net phosphorus quotients for the same time periods by factors ranging from two to twenty. The absolute quantities of nitrogen and phOSphoms fixed by the community were nearly the same (table 15). Therefore, the difference in the net TABLE 13 NI TRJGEN/NEIGHT QUQTIENTS OF PERIPHYTON COLLECTED FROM ARTIFICIAL SUBS'IRATA Collection Mean N Mean w Mean N: 103 Standard Error Date (mg) (mg) 3m N (mg) W.(mg) 1958 (3:) (_+_) 5 May 2.18 1111.60 15.110 0.283 7.01 12 May 1.09 112.110 9.70 0.071 6.01 19 May 1.19 53.07 22.112 0.191 2.82 11 June 2.110 1112.87 16.80 0.1111 11.115 18 June 2.111 207.90 11.59 0.1111 36.51 25 June 2.08 223.17 9.32 0.233 72.15 TABLE 111 NITROGEN EFFICIENCY QUOTIENTS CALCULATED FOR 1001NETERS OF RIVER Collection Exposure % of Total Net Date Period Inorganic Efficiency (Days) N Import <1 Quotient (i) 1958 5 May 10 1. 51 0.027 12 May 7 0.811 0. 0211 19 May 7 0.28 0.080 291%ay 10 0.18 0.131 11 June 9 0.73 0.062 18 June 7 1.h3 0.032 25 June 7 0.21 0.187 total inorganic N import computed from 27 November 1957 to TABLE 15 NI'I‘ROGEN/PI-KEPHORUS QUOTIENTS OF PERIPHY'IUN COLLECTED FRQI ARTIFICIAL SUBSTRATA Collection Mean N Mean P Mean N Standard Error Date (mg) (mg) m N (mg) P (mg) 1958 (1) (1) 5 May 2.18 2.12 1.03 0.283 0.1633 12 May 1.09 1.72 0.63 0.071 0.13511 19 May 1.19 0.97 1.23 0.191 0.0997 29 May 1.29 0.88 1.117 0.099 0.1003 11 June 2.110 1086 1.29 0.1141 0 18 June 2.111 3.76 0.611 0.1111 0.02011 25 June 2.08 1.70 1.22 0.233 0.2507 83 811 efficiencies were due to the greater evailablity of inorganic nitrogen. During the study period the ratio of inorganic dtrogen to soluble phos- was approximately five to one. The relation between the phosphorus and nitrogen content of peri- phyton was studied from 5 May to 25 June 1958. Nitrogen/phosphorus quotients for this period varied from 0.63 to 1.197 with a grand mean of 1.011 (table 15 ). Working with laboratory cultures of several Species of green algae and diatoms, Ketchum and Redfield (19119) noted N/P values ranging from 1.6 to 5.3. Correll (1958) found that diatoms growing in an oligotmphic stream had N/P ratios of about 10. These data and information from the Red Cedar seem to indicate that N/P values are dependent upon the avail- ability of phosphorus and the growth rate of the algae. That is, algae grown in phoephoms rich media tend to have lower ratios of N to P because of phosphorus storage. In addition, growth inhibition favors even lower N/P quotients because of the associated accumulation of polyphosphates. Pollution 85 86 The biological evaluation of stream pollution is based primrily on taxonomic changes in the biota. The presence or absence of certain indi- cator species along with their relative abundance has long been used as a criterion of pollution (Turner 1927, Gaufin and Tarzwelli 1952). Such methods are, at best, semi-quantitative and often deal; with organisms whose ifllentification is difficult and uncertain. Patrick (19119) Save a- quantified system for evaluating pollution based on the taxonomic compo- sition of the bionase. Patrick's work is significant because it extends the concept of indicator species to indicator communities and shows the inadequacy of chemical analyses for pollution detection. However, this method cannot be seriously regarded as a- field technique by pollution biologists because of the time and taxonomic knowledge required. A functional rather than a taxonomic approach to pollution evaluation has been suggested by Odum (1956). Odum classified lotic comunities as being oligosaprobic, mesosaprobic, or polyeaprobic on the basis of the ratio of comunity oxygen production to community respiration. This is essentially a quantified version of earlier attempts to categorize pollution by the extent of oxygen depletion. However, unlike previous methods it provides a continuum of irxiex figures (production/respiration quotients) ranging from oligosaprobic to polysaprobic communities. These figures are easily obtained from standard D.O. determinations made in the field. Unfortunately, such a classification is applicable only in cases of organic pollution. Data from the Red Cedar River indicate that the functional composition and physiology of periphyton colonizing artificial substrata might be useful indices of pollution. ON and P/N quotients. The floral components of a periphyton colornr can be placed into two functional groups; the producers, and the reducers. 87 In a clean-waster zone the conmumity is composed primarily of autotrophic organisms. In comtmt, extreme organic pollution is characterized by a large proportion of heterotrophic plants. Therefore, in the tramition from oligosaprobe to polysaprobe there should be a continuum of index figures, based on changes in the proportion of producers to reducers, which reflect the degree of organic pollution. The relative proportion of producers to reducers in periphyton colonizing artificial substrate— was studied using the following quotient. C/W x 1,000 where C - the corrected absorbancy of phytopigments extracted from the colorw. w - the ash-free dry weight (mg) of the celery. This quotient makes use of the fact that a- quantitative estimate of the weight of a diatom pepulation can be made from the extracted phytopigments (page 3h). Therefore, C is proportional to the weight of the producer fraction of the colony. Ae the proportion of reducers in the colorw becomes greater W will increase and the ratio of C to W becomes smaller. C/W ratios obtained from a clean-water zone, a polluted zone, and a recovery zone are shown in table 16. The polluted zone was sampled at a point 25 meters downstream from the sewage outfall of the Williamston treatment plant. The clean-water site was located at Dobie Road.and the recovery zone 50 meters downstream from the sewage outfall. ‘ The difference in quotients between the polluted zone and the clean- water zone are quite distinct. Mean C/W ratios from the now-polluted zone varied from 10.02 to 16.91. Mn values obtained from samples 25 meters downstream from the sewage outfall were 2.81; and 1.81;. A‘ mean 88 c/w quotient obtained 50 meters downstream from the sewage outfall was essentially the same as one obtained from the clean-water zone on the same day (16.91 vs. 18.89). This dramatically demonstrates the rapid recovery of the stream to an oligosaprobic state. Brehmer (1956) found that a septic zone was never famed, even in the inrnediate vicinity of the sewage treatment plant. 0n the basis of Odum's‘ classification (1956) the intensity of pollution was, at worst, mesosaprobic. Therefore, it was not possible to obtain C/N quotients from the full gamut of saprobe states. Nevertheless, it was demonstrated that CAI ratios can be used to differentiate between oligosaprobe and mild mesosaprobe. It seems logical to assume that polysaprobic communi- ties would have their own characteristic C/W value. When considering extreme polysaprobic conditions, such as described by Bartsch (19118), where extensive stands of "sewage fungus" exist, one can be reasonably certain that the ratio of producers to reducers will be very small. Thus far only the evaluation of pollution from sanitary wastes has been considered. Date from the Red Cedar indicate that periphyton tech- niques might have some limited value in detecting sublethal doses of industrial pollution. Since the degree of industrial pollution present in the Red Cedar is unknown the following discussion is more Speculative than factual. It has been demonstrated that algal growth inhibition results in interred. polyphosphate accumulation (Nihei 1955). Therefore, any compound which inhibits algal growth would cause an increase of phosphorus within the periphyton colony. Such a starts would be reflected in the. quotient of the phosphorus content to colony weight, or in the analogous ratio of phOSphorus content to the absorbancy of phytopigments extracted TABLE 16 A COMPARISON OF C/W QUOTTENTS OBTAINED FROM A CLEAN-WATER ZONE, A RECOVERY ZONE AND A POLLUTED ZONE Collection C/W Quotient Date Clean-Water Recovery Polluted 1957 1 Oct 16.91 18.89 18 Oct 10.56 ‘ 1.8h ’4 NOV 10e02 2.81; 89 from the colony. It hes been shown.by Procter (1958) and.IawTence (1958) that a wide variety of compounds, both inorganic and organic, can inhibit algal growth. Such an index of pollution is subject to severe limitations. Date from the Red Cedar (page 72 ) and elsewhere (Lund 191:9) have shown that growth inhibition and subsequent polyphosphate accumulation is a natural occurence in non-polluted waters. It is also unlikely that PIN or P/C quotients occur in a quantitative continuum because of the many factors (i.e. species composition, nutrient supply, growth rate) that contribute to their magnitude. Therefore, the use of such quotients would have to be on arcomparetive basis (cleanewater‘vs. suspected pollution) and be limited to detecting rather than evaluating pollution. Besides its sim- plicity, the only advantage this technique might have is the capability of detecting extremely low levels of pollution. SIMMARY 91 92 1. At base flow the Red Cedar River has a methyl orange alkalinity of 250 ppm and.a pH of 8.1. The amount of normal carbonate present throughout the year is negligible. The total solids content of the stream varies between approximately 200 and 300 ppm. 2. Suspended solids act as a selective filter favoring the trans- mission of longdwaved radiation. At all levels of turbidity red trans- mission exeeeds green, and green transmission exceeds blue. Dissolved color fluctuates independently of turbidity and plays only a minor role in influencing the total quantity of light transmitted to the stream bottom. 3. The absorbancy of extracted phytOpigments can be used to make quantitative estimates of periphyton weight if corrected to conform with the Lambert-Beer Law. Productivity measurements based on phytopigment units showed excellent agreement with gravimetric productivity estimates. Successful measurement of primary production by diurnal oxygen curves appears to be limited to periods of intermediate productivity. h. The seasonal range in net primary production as measured by artificial substrata is 0.01 to 2.28 g m"2 day-1 with an annual mean -2 _ of 0.56 g m day 1. The annual gross rate of periphyton production is 2 day-1. The preceeding rates are given in estimated to about 1 g m- terms of ash-free dry weight. 5. Photosynthetic efficiencies based on net production and surface radiation vary from 0.003% to 0.2h5% with an annual mean of 0.07%. The annual.mean efficiency based on gross primary production and surface radiation is estimated to be about 0.1%. The annual mean rate of energy fixation by periphyton is 0.23 g cal cm'2 day-1. This figure represents the amount of energy available for transfer to consumer organisms. 93 6. The annual import of phosphorus into the study area is 16 metric tons. During three flood periods collectively representing a tine lapse of 30 days about h5% of the total phosphorus import for the yeat was carried into the study area. 7. Phosphorus fixation by periphyton is closely related to peri- phyton growth. When the two rates are plotted, the relationship resembles an exponential curve. The ratio of phosphorus content to the ash-free dry weight of the periphyton colony (x 1000) varies from 1.5L to 33.63. In general the slowest growing colonies have the highest phosphorus con- tent. Seasonal phosphorus efficiency quotients based on total phosphorus import and calculated for a 100 meter length of stream vary from 0.0003 to 0.hl%. Similar efficiency quotients based on soluble phosphorus range from 0.0007 to 0.61%. 8. The total inorganic nitrogen import for a nine month period was 8h metric tons. The effect of floods on the seasonal distribution of in- organic nitrogen was less pronounced than in the case of phosphorus. The data appear to indicate that runoff is the primary source of inorganic nitrogen. 9. The nitrogen content of periphyton samples was l.h9% of the ash-free dry weight. The ratio of nitrogen to phosphorus in periphyton samples collected during periods of maximal growth is approximately 1. Inorganic nitrogen efficiency quotients calculated for a 100 meter length of stream range from 0.03 to 0.19%. 10. The ratio of phytopigment absorbancy to periphyton colony weight can be used to differentiate between oligosaprobic and mesosaprobic com- munities. Such ratios obtained from a clean-water zone fall between 10.0 and 16.9. Similar values obtained from a polluted zone vary from 1.8h 9b to 2.9. Ratios based on the phosphorus content of the colony are less reliable indices of pollution and appear to be limited to pollution de- tection rather than evaluation. LITERATURE CITED 95 96 Alexander, G.R. 1956. The fertilization of a marl lake. Unpub.!flaster's thesis,1flich. State Univ. 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