Closed Ecological Systems Author(s): Frieda B. Taub Source: Annual Review of Ecology and Systematics, Vol. 5 (1974), pp.

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Copyright 1974.All rightsreserved

CLOSED ECOLOGICAL 1,2 SYSTEMS

Frieda B. Taub
of Washington, 98195 Collegeof Fisheries,University Seattle,Washington

.4073

INTRODUCTION Sealedecologicalsystemshave been surprisingly unexplored as ecologicaltools. Thereis a fascination in exploring thebehavior of systemsfor whichall the interacting partsare accountable. Althoughthe "balanced aquarium" was once said to be of seriousstudiescouldbe located.The investigalittle documentation fashionable, tion of bioregenerating life-support systemsfor space travel enjoyedonly a brief of the hydrogen-oxygen fuel cell severely flurryof activitybeforethe development limitedtheirpotentialusefulness. Somemicrocosm and continuous cultureexperiments,althoughnot sealed,permitinputand outputmeasurements and therefore of total systems.These approaches approachrepresentation can be used to test hypotheses concerning the homeostatic regulating mechanisms postulated to exist in natural communities. Thepastworkis reviewed herefor the convenience of those this area of research.The U.S. space-related considering studies are emphasized since this literature has not come to the attentionof many biologists. Althoughthe phrase"closedecologicalsystems"has gainedpopularusage,the i.e. lightinput arealmostalwaysopento energyexchange, systemsunderdiscussion and heat loss. Were this exchangenot permitted,e.g. a bacterialculture in an insulated bottle,the functioning of the systemswouldbe limitedby theinitialsupply of organic of temperature. energyandby theelevation Spacevehiclesmayhavetheir own energystores,but they dissipateheat. In fact, most of the regenerative lifeare far frombeingclosed even to materials. Someaquariato supportexperiments whichfood is not added,but whichexchangegas with the atmosphere, are spoken of as "closed"; most microcosm yet C, 0, H, and N exchangegaseously.However, in this manner, havebeenconducted on the premise thattheyaremore experiments realisticthan totallyclosedsystemsbecausenaturalsystemsare opento the atmosphere.
'Contribution No. 396, Collegeof Fisheries,University of Washington. 'This workwas partially supported by the U.S. Environmental Protection Agencyand by the National ScienceFoundation underGrantNo. GB-20963to the Coniferous ForestBiome, EcosystemAnalysisStudies,International BiologicalProgram, Contribution No. 111. 139

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REGENERATIVE LIFE-SUPPORT SYSTEMS FOR SPACE FLIGHT a life-support systemthat will supplyman'sneedsMannedspaceflightrequires H20, and food-and eliminatehis wastes-CO2, urine,feces,and heat. Early and storage of water, food, and systems relied on expendableCO2absorbants pressurized oxygen;thus the weightof the storedmaterialsrose sharplywith the durationof the mission (54). The attractiveness of convertingwastes to needs of potentialregenerative systems.For long, multimanned prompted an evaluation and complexities of suchsystemswould missions,the weight,powerrequirements, materials. In the be less thanthoseof the weightandstoragesystemsof expendable was mid-1950sno one couldpredictwhenthe needwouldbe critical,but research classes: systemsconsidered wereof threemajor begunenthusiastically. Regenerative (Hydrogenomonas and(c) chemosynthetic (a) physicochemical, (b) photosynthetic, bacteria).
02,

Physicochemical RegenerativeSystems fromsimplefiltration of urineandcabincondensate anddistillation Systems ranging of H20 of potableH20 to moreelaborate for regeneration systems,e.g. electrolysis reactionfor recovery for recoveryof 02 (H20 -4 H2 + 1/2 02), to the Sabatier of methaneand water (CO2+ 4H2 CH4 + 2H20) coupled with crackingof methane(CH4 + H20 -4 3H2 + CO) for recoveryof the H2 to maintainthe human andpotentially to synthesis via polymerization to produce Sabatier reaction, food compoundsfrom methanoland formaldehyde (25). Most physicochemical but were amenableto an engineering apmethodshad high energyrequirements proach.Techniquesof water recoveryhave been evaluatedand compared(95). on research interestsincethe concurrent Theseeffortsare now largelyof historical batteries. energystorageled to the use of the H2-02 fuel cell in placeof the heavier of the fuel cell madethe human02 requirement seem petty, The 02 requirements To some, andits wasteproduct of waterwasfarmorethanthe humanrequirement. the use of a waste productof the fuel cell to supply a humanneed would be use.Various considered physicomultiple, nonregenerative cycling,butit is actually chemicalatmospheric controlsubsystems were proposed(2), and a recentsystem in a 90-daytest (35). was appraised PhotosyntheticRegenerativeSystems A multiplicityof systemswas rapidlysuggested.Initiallythey werejustifiedas as food was also considbut the potential use of plantmaterial C02/02 converters, ered.Many weredesignedalso to utilize urineand wash waterand possiblyfeces to supply the algal nutrients.It was generallyacceptedthat a three-component human would be requiredas a minimum; system (man-wasteprocessing-plant) nutrients for wastesnot directlysuitable wouldbe processed to supplythe required algal growth. Various short circuits and alternatepathways (Figure 1) were proposed. The earliestplanfora photosynthetic life-support systemis not knownto me,but the requirements it must havebeendeveloped before1954,whenMyerspublished

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CLOSEDECOLOGICAL SYSTEMS
MULTISTAGE ECOLOGICAL SYSTEM

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Secondary Consumer

WASTE
STORAGE

Primary Consumer

HCE1

GreenA Plants

FigureI A compositemultistage life-support system.

for a man-daysystem.Its possibility had been suggestedin 1953by Bowman(8). By 1956,a conference on photosynthetic gasexchangers wassponsored by the Office of NavalResearch (93);systemsweredesigned to prolongthe submergence capability of nuclearsubmarines withoutthe releaseof gasbubbles. In 1959Rich et al (76) presenteda fairly advancedscheme,with water reclamation, photosynthetic gas exchange,and wastestabilization. Tischer(87) discussedvarioustypesof systems: human-yeast-bacteria-algae; higherplant-algae-waste-human; the aboveplus an animal-goat, Daphnia, or slug(?); hydroponically grownplantson a bacterial-algal converter;a Hungatecycle, crew-microbes-plants; and a van Niel microcycle, Artemia, bacteria, and algae.None of the proposed systemscould meet all human needsnor recycleall productsand they werelargelyprojections of what could be done. The parallelbetweenthe earth'secologicalcycleand the proposed spacesystems wasimmediately obviousandwasinfluential in theirinitialdesign.Surprisingly, few ecologistsbecameinvolved,as is discussed later.Development wentlargelyto civil engineers (sewagetreatment processes), plantphysiologists (massalgalcultureand higherplants),medical andhumanphysiologists (humanrequirements), andchemical engineers(scaleupof sewageand algal systems). Although all studies claimedto be directedtoward the development of total systems,most concentrated on the processeswithina single subsystem. Artificial inputs were often used to standardize experimental conditions,e.g. an air-CO2 mixturefrom a pressurized tank ratherthan fromhumanrespiration. Usuallythe

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outputwasmeasured butnot usedby another Thisuncoupling of subsyssubsystem. tems,whilemakingthe conductof eachprojecteasierand independent, eliminated or the effectsof tracematerials. feedbacks, interactions, The studiesreviewed here areorganized as singlesubsystems, coupledsubsystems, and the holisticapproach. The clashbetweenthe subsystem and holisticapproach is discussed as an introduction to the latter.
SINGLE SUBSYSTEMS

Human Daily Balance An enormousamountof medicaland physiological researchwas reviewedand performed to determine humanrequirements and waste productsunderthe likelystressesof spacetravel.Most of the pertinent data were Data Book (97). The workis a convenient broughttogetherin the Bioastronautics document of humanfactorvalues.Further are nutritional andwasteconsiderations compiled in (57). The nominal values for regeneration studies are shown in Table 1 (1). Algal Culture Algal oxygenor cell production receivedthe overwhelming effort of thenonmedical studies. in 1966thatabout$30millionhadbeen Jenkins estimated spent on researchinto photosynthetic bioregenerative systemsduringthe 12-15 in 1953:(a) preceding years(37). It was sparked and directed by two publications Earlierstudies on mass algal culturewere made conveniently availablein Algal Culture to PilotPlant (11), a compilation from Laboratory designedto encourage consideration of masscultures as a sourceof high proteinfood to solvethe world's foodproblem. TX 71105,a high temperature (b) Chlorella pyrenoidosa strain,was defined as the mostefficient andits growthandproduction photosynthetic organism, ratesweredescribed that photosynthetic (78). Myerssoon estimated gas exchange fora man-day wouldnecessitate in 230 litersof liquid 2.3 kg of Chlorella suspended 10 hp (52). The high powerrequirehaving24 m2of lightedsurfaceand requiring of 19%for conversion of electricity to fluorescent mentwas basedon an efficiency light and 10% for conversionof light to photosynthesis. The high volumeand of densealgal surfacearearequirements werenecessary becauseof the self-shading A significant as well as number of aviation anduniversities, suspensions. companies the U.S. Air Force,initiated a genstudiesalmostsimultaneously. Thereprevailed
Table 1 Nominal values of the daily materials balance for man in space (1). (Values by other authors are not materially different.) In Total: 3615 g Total: Water: Out 3615 g 2730 g 840 g (19.2 moles)

Water: 2400 g Oxygen: 715 g (22.3 moles) Dry food: 500 g (95% digestible) (2400 cal, of which protein = 10%, fat = 35%, and carbohydrate = 55%)

Carbon dioxide: Dry solids: 45 g

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SYSTEMS CLOSEDECOLOGICAL

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wouldprovecrucialto the spaceeffortand eraloptimismthat this line of research was that the and a generalsupposition earthapplications, might have important eitherthe U.S. Air Forceor NASA, wouldsupportthe more federalgovernment, teams. promising amassedon the cultureof algae, especiallyof ChlorellaTX The information systems (29 in number)were reviewedand Significant 71105, is overwhelming. for one-mansupportvariedfrom 3.5 to in 1966(49). The requirements compared 3,000 liters, from 2.5 to 71 m2, and from 7 to 100 kw. In most instancesdense high surfaceareasbut small volumeswere tradedoff for those culturesrequiring but less dense requiringless surfaceareas, becauseof deeperlight penetration, greatervolumes.Most of those systemswould requiremultiplesor scalingup to achieveone-mansupport. to NASA;especially In spiteof thiswork,thealgalsystemsdid not lookfavorable Althoughas muchwasknownabout of reliability. werethe unknowns objectionable and their as any cell, therewas no hope of knowingall its components Chlorella underall conditions.Becauseof the inefficient separateand combinedproperties inefficient to light(lessthan20%)andthe subsequent energy of electrical conversion no more than 20%), at best effi(probably conversionof light to photosynthesis as compro4%. The use of sunlightwas rejected ciencywouldbe an unacceptable to the constantorientation of the spacevehicleshell, requiring misingthe integrity events. and other unpredictable to sunbursts sun and beingvulnerable for Higher Plants Becauseof the potentialadvantagesof lower requirements as lowerweightof the system,and the morereadyacceptance water,and therefore photosynon higherplantsas potential humanfood, a limitedeffortwas expended stages. thetic agents.None of this effortwent beyondthe preliminary minimum demonstrated Duckweed(severalspeciesof the family Lemnaceae) aldivisionrates rangingfrom 12-20 hr (vs 8 hr for thermophilic Chlorella), though the authorscautionedthat light intensityand other conditionswere not directly is thatgas wasexchanged of the duckweed A majoradvantage comparable. so that elaborategassingand degassing betweenthe plant and the atmosphere, the plantsweresepawas also simpler; Harvesting werenot necessary. procedures (96, 98). Otheraviation ratedby coarsescreensanddid not haveto be centrifuged companies,such as RepublicAviation(75) and the MartinCompany(32), were growinghigher plants, but their detailed reportswere not in my collection of literature. fieldsweretestedon and gravitation The potentialeffectsof alteredatmosphere werenoted,butthe resultswerepreliminary snapbeanandotherplants.Differences of gravitational on studiesof higherplantsaregivenin (4). Studies (4). Otherreports but on the Biosatellite program, of plantgrowthhavecontinued effectson patterns effort. as a basic research studiedas a WasteProcessing Biologicalwaste processingwas more frequently and the materialis reviewed partof a coupledsystemthan as a singlesubsystem, in that section.Whenwasteproductswerenot recycledthey couldbe storedwith releasedfrom the vehicle,or incinerated. a disinfectant,

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COUPLED SUBSYSTEMS

Photosynthetic-AnimalRespiration Even coupled photosynthetic subsystems were

that Myersperformed notablyfood. It is probable transfer, usuallyopento material between1955and 1958cited in (9), but I have the initialalga-mouseexperiments not seen the Air Force Reports. nidulans a 15litercultureof Anacystis reported The Schoolof AviationMedicine subsequently and 80 for hr, mouse g 30 one with was coupled alga) (a blue-green with three 30 g mice for 80 hr (31). In each instance,the abilityof the cultureto soonexceeded andthe 02 concentration the animalrespiration 02 exceeded produce was high at the startand was not 30%by volume,whereasthe CO2concentration in the CO level, increase findingwas a continuous depleted.The most troublesome of a COburner by inclusion was eliminated the problem to the Anacystis; attributed (also a blue-green (Hocalite,heatedto 1250C).A 15 liter cultureof Synechocystis of four mice. In another alga) was similarlyable to match the 02 requirements cultureunit maintained the algal in not reported but mentioned experiment detail, only gas exchangewas coupled.The mice for 15 days. In all of theseexperiments lab chow,crumbsof whichbecamemixedwith fecesand micewerefed a standard mediumand defined urine.The algalculturesweregrownentirelyon a chemically mice. of the waste products the gaseous wascoupledwitha 3 kg primate Whenthe 15.5literalgalunitwith Synechocystis was terminated it suppliedonly one sixth of the oxygenneeds,and the experiment hr (30). at 21/2 before1960to demonstrate At ChanceVoughtAircraft,studieswereunderway of mice can supportthe 02 requirements strainof Chlorella that the thermophilic becauseof excessiveCO2and threewere (8). Of eight trials,five were terminated and wereendedon the 12th, 14th,and 28th day for mechansuccessful considered 45 days Oneof the micetolerated functions. to thebiological unrelated icalproblems with 10% at one time an atmosphere experiencing atmosphere, in the regenerated the respiratory and 10%02. Onthe longestruna 4 literalgalculturesupported CO2 from 21% to a maximumof 30%. needsof a 30 g mouse,and oxygen increased 20 and31% between wascontrolled Whentwo micewereusedthe 02 concentration 02 was observedto drop slowly to 10% afterday 7 and until day 15, whereEfter towardthe end The imbalance was terminated. until day 20 when the experiment fecesand wettingof the accumulated to the accidental wasrelated of the experiment Clearly, bacteria. the by and uptake 02 increased activity urine and subsequent supportto the mice, it could not althoughthis systemcould providerespiratory waste products.The aerobic supplythe biological02 demandof the accumulated with the wasteswas equalto two mice. Methaneand ethane associated metabolism a 40 g mousewaskeptin goodhealthfor 66 days; werenoted.In a laterexperiment the 02 increasedto 63% and CO2was usually controlledat 0.2% or less. No methane,ethane,or CO were found (9). ellipsoidea studiesbetweena dwarfmouseand Chlorella gasexchange Additional between imbalances fromslightbutpersistent in excess02 production againresulted the C02/02 ratiosof the mouseand alga (27). These ratioscan be shiftedby the

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of the mousediet and the formof nitrogen(nitrateor urea)supplied composition to thealga.Metabolism of wasteproducts wasminimized by drystorage. Twosealed experiments of 15 and 24 days were terminated, not becauseof any component failure,but becausean increase in oxygenand total gas volumeexceededthe limits of a variablevolumecomponent. They suggestthat earlierexperiments by others probably had significant gas leaks. The Bioregenerating Unit (BRU) was a morecomplexsystembut also restricted to gaseousbalance.It included culturesystemsof the fungusLinderina pennespora, the high temperature strainof Chlorella pyrenoidosa, a sewagereactor,and a rat (45). The funguswas selectedfor its abilityto convertgaseousammonia to protein, a potential animalfeedsupplement. The 2.5 literalgalculturewas supplied with all its requirements in its growthmediumexceptcarbon,the fungalculturewas suppliedwith all of its growthrequirements in its medium,and the rat was provided with food (unspecified, presumably a standardlaboratory chow) and water.The feces and urineof the rat werestored,but gas productssuch as NH3 would have circulatedthroughthe air and have been availablefor fungalincorporation. The sewagereactorwas runon rat and rabbitfecescollectedfromotheranimals.Thus the systemtestedthe abilityof the algalcultureto provide02 forthe fungus,sewage reactor, and rat. Gas balancewas achievedfor 48 hr on a trialrunwithoutthe rat. Failureresultedafter22 hr with the inclusionof a 65 g rat and sewageload. Until thena reasonable 02 balance hadobtained, butCO2hadincreased from0.08 rapidly at 4 hr to 1.20%.By 28 hr the 02 haddeclinedto 8.4%and the CO2had increased to 10%.Thesetrendscontinued until the rat died, at approximately 35 hr. By 72 hr the sedimentable sewagesolidshadbeenreduced75%, and by 92 hr the sewage had been digested.London& West (45) attributed the gas imbalance to a sudden 02 demandby the fungalor sewagesystemand not to a malfunction of the algal system.Theynoted,however, that the algalcountbeganto peakoff at 40 hr, as in their earlierexperiments. An improved BRU had9 litersof algalculturewith increased lightandcontinuous mediuminputand culturewashover. The systemwas runfor 174hr with CO2 gas to primethe algalculture; thena 149g rat,feces,and fungalsporeswereadded and the systemwas sealed.After24 hr the CO2had risensharplyto 7%, whereas the 02 had dropped I. At 70 hr algalculturewas shiftedfromcontinuous to 11% to batchoperation, but gas imbalance continued to worsen.Duringthis timeof severe stressthe rat inspired deeplyand rapidlybut continuedto takefood and water.At 80 hr, half (1.2 liters) of the fungalculturewas removedin orderto lessen the 02 demand; a yieldof 3 g dryweightwasrecovered. Thereafter gasbalance returned; the 02 increased to 17%andthe CO2fell to 2.6%.Additional mediumwassupplied to the algal cultureduringthis time. At 216 hr the run was discontinued because of mechanical breakdown. The rat appeared to be healthyand had gainedweight. A subsequent runof 309 hr resulted in a heavyvacuumand was thendiscontinued. The seriesof experiments was interpreted as showingthe possibility of photosynthetic regeneration of the atmosphere. Additionalexperiments indicated that bacterialcontamination of the algalculturewas not responsible for the problems. After a total of 819 hr (34 days), the algal culture was still vigorousand capableof

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Fungalgrowthoccurredapparits role as a gas exchangemechanism. performing ently withoutthe use of gaseousNH3,the cornsteep liquorin the mediumhaving edible,was not supplyof nitrogen.The fungus,presumably providedan adequate tested. pyrenoidosa by a 57 liter Chlorella A 1220 g Cebus monkeywas maintained culturefor several50 hr periods(99). The 02 graduallyrose from 21 to approxiactivitywasthe major leveldidnot exceed1%.Themonkey's mately25%.TheCO2 was The02 production in 02 andCO2concentrations. fluctuations causeof observed nutrientexchange,and controlledby light intensity,nutrientfeed concentration, CO2supply. controlfor humansubjects,. Algalcultureswerealso usedto provideatmospheric made the decisionto bypassanimalstudiesand go BoeingCompanyresearchers of 55 liters systems(7). Multipletankswith capacities directlyto man-supporting eight wereeventuallyused. Like Golueke& Oswald(33), each were constructed; on 02 productionper unit volume and later came to the they first concentrated perunitareaandlight intensityaremoreimportant. that 02 production conclusion peaked24 hr afterseedingat 43 trials02 production unmanned In the preliminary ml 02/liter hr, or 2.4 liters 02/tank hr. It was estimatedthat eight tanks would or low-activity of one manin a resting provide19 liters/hr,or aboutthe requirement but no contaminated, The algalcultureswereknownto be bacterially environment. the completed: effectswereseen. A 56 hr mannedrun was successfully deleterious 02 remainedbetween 19 and 21% while the CO2 remainedbelow 2%; minor activityand the periodicharvestof wererelatedto the humansubjects' variations that a man studieswith smalleralgaltanksindicated the algalcultures.Subsequent on much less than the 380 liters used. could be supported Algal-AnimalFeedingStudies The literatureon algal-animalfeedingstudiesis (43, 48), but the resultsare considerable and much of it has been summarized was highly that Chlorella For instance,Lubitz(46) reported conflicting. frequently the sole sourceof proteinin an well absorbed, and capableof providing palatable, otherwisecompletediet for the rat. Conrad& Johnson(16) had poor resultsand et al (94) wouldbe poorlydigestedby the rat. Vanderveen assumedthat Chlorella for some toxic effects mightbe responsible contamination suggestedthat bacterial reportedin humanfeedings. in resultswithinthe animalexperidifferences I suggestthatsomeof the extreme was when Chlorella feedingtechniques; mentscouldhavebeendue to the different but whenprovidedas the used as a portionof a completediet it was satisfactory, aboutthe lackof calcium I amparticularly concerned soledietit wasunsatisfactory. in most algal growth media; Chlorellaapparentlydoes not requirecalcium in but ratsdo. In the humanfeedingtrialscontrolovertotal diet substantial amounts, The generalconsensusis that algae, and particularly was sometimesinadequate. may not be suitableas humanfood;but no toxins have been identified. Chlorella, to be an open question;Lubitz(46) was the only one to I regardthe digestibility Otherssimplyquoteearlierworkers good assimilation. measure it, and he reported as statingthat the wall is resistantto digestion.

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Human WasteProducts to Algal Nutrients The direct or indirectuse of human forphotosynthetic wasteproducts in mostschemes, recycling wasassumed although additional supplements mightbe deemednecessary. Eachmethodof treatment had its own advantages and disadvantages (72). The directutilization of wasteswouldrequire a combined aerobic bacterial-algal reaction chamber. Pipes(72)statedthatthe algaewouldalwaysproduce 02 in excess of that required for the oxidationof the organicmatterif light and nutritionwere adequate.The criticalproblemwould be the concentration of CO2availablefor photosynthesis. Sucha systemwasestimated to require a detention periodof 60-72 hr and a capacityof 30 litersper person.It wouldbe greaterin size than the unit required for the otherprocesses, but wouldallow othercomponents of the system to be correspondingly diminished. The advantages of such a unit weresaid to be: 95% oxidationof organicmatter,recoveryof potablewaterafterdisinfection and other treatment,and production of a food supplement consistingof a mixtureof algae,bacteria, and a few higherinvertebrates. The furthestdevelfungi,protozoa, opmentof such a system was the "Algatrons" (34), 11 of which, each 18 in. in diameterand 4 ft long, wouldbe required per man for gas exchange,wastetreatment, and waterrecovery.In experiments, Oscillatoria (a filamentous blue-green alga) gave 02 yields comparable but havingan unpleasant, to those of Chlorella, odor.The unitwasalsosubstituted swampy intothe "microterella," discussed later. A manned of the mouseunitwasdesigned equivalent termedMECCA(Man-AlgalBacterialEcologicalCultureChamber). Directutilization of wasteshadobviousdisadvantages: diseasetransmission, light anda strongpsychological revulsion absorption, to the potential astronauts. Numerous methods were suggestedby Pipes (72) to overcomethese: incineration, wet oxidation,chemicaloxidation,aerobicmicrobialdigestion(activated sludge),and anaerobic digestion.An electrolytic methodof breaking down "dailyoutput"was developed,and the products,combinedwith additionalurineas a supplementary nitrogensource, supportedgrowthcomparable to the standardKnop's medium (50). In furtherexperimentation micronutrient and reliability supplementation betweenbatchesweretested(51).Theproducts in verylimited of combustion resulted growth unless additionalCO2was supplied;with CO2 and the chelatingagent EDTA (ethylenediamine tetraacetic acid) growthyields were 107 and 108cells/ml
(74).

Themicrobial treatment of wastes,separated membrane by a semipermeable from an algal culture,was tried(6). Experiments over periodsof 21 daysdemonstrated successful The system algalgrowthon the diffused productsof bacterial oxidation. was not entirelyfunctional froman engineering pointof view becauseof the slownessof waterdiffusion. Bieberdorf (6) notedthat,duringa 20-dayrun,not only was of waste, goodgrowthof algaeobtained accompanied by satisfactory decomposition but the composition of air withinthe chamberremained constantat 20% 02 and 80% N2 even though it was sealedfrom the laboratory It may also atmosphere. be worth noting that holes developedin the cellophanemembranes in 4-5 days when one membraneseparatedalgae and fungi but not when two membranes wereusedseparated by a barrier of distilledwater.In the actualexperiments, syn-

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thetic wastes consistingof dehydrated cooked meat, distilled water,and papain were used. As a test of mesophilic activated sludge,cellulose,soaps,detergents, andvomitus wereeach added.Cellulosewas degraded afteran initiallag (67). Castilesoap, the sodiumdodecylsulfate,andvomitushadno deleterious detergent effects, but nonbiodegradable detergents were a problem(68). Human Waste Products to Potential Food It wassoon recognized thatsomeof the humanwasteproducts mightbe usedin producing an alternate foodsource.Tischer & Tischer(89) studiedthe use of humanurineand feces for the production of the crustaceanDaphnia, the mold Rhizopusdelemar (alone or combinedwith the bacterium Pseudomonas), the protozoan andthe yeastSaccharomyces Paramecium, cerevisiae. EarlierTischer(88) had shownthat R. delemarconvertsup to 50% of fecal nitrogeninto fungalprotoplasm. Fecal nitrogenwas not as good a sourceof for fungalgrowthas nitratenitrogen. nitrogen Improved yieldsof moldgrowthand efficiencies of conversion of the feces with mightbe obtainedby supplementation certainformsof nitrogen,carbon,and possiblyvitamins.Mold grownon human wastesubstrates was driedand fed to fivehamsters as a sole food for 10 days and was observed to produceno weightchange.This resultwas interpreted as showing that molds cultivatedon humanwastescan be convertedinto animaltissue. At the Universityof Washington studiesweremade that involvedfeedingfish, Tilapia,on algae and intermediate zooplankton grownon humanwasteproducts (81, 82). Threetypesof 17-liter systemsweretested:(a) a feces-supported communityof algae,Daphnia,rotifers, and Tilapia, protozoa, (b) a similar community that was unfed,and (c) anotherthat was fed frozenbrineshrimp.They were aerated duringthe 93-dayrun so that temporary would not be critical.At gas imbalance the end of 30 days the fish were weighedand replaced.The fish in the unfed wereobviously whereas ecosystems starving, thosein the ecosystemreceiving feces hadgrownalmosthalfas muchas thosein the ecosystem fed brineshrimp. Thereafstarved ter, two systemswerefed feces and the thirdbrineshrimp.The originally fishgrewbut nevercaughtup withthe othergroupsin average or totalweight.The fishin the ecosystem thathadreceived fecesfromthe beginning wereapproximately of the sameaverageweightas the controls,but fewersurvived.Mostmortalities in the systemsweredueto theentrapment of fishin tubesintended to shelterzooplankwerehigherin the feces-fed ton, but mortalities systems.Althoughit was intended that the fecal materialbe utilizedas a nutrientfor the algae and microbes,which wouldbe eatenby the invertebrates, and these,in turn,wouldbe eatenby the fish, we observedinsteadthat the fish activelyfed on the fecal materialas it was being added. Gas analysisof the inputand outputair indicateda balancein the systemafter the firstfew days. No unpleasant odors developed.The Daphnia,in spite of the attemptto shelterthem, were all eaten within the first few days. The Chlorella underwent a successionof bloomsand then steadilydiminished.Ankistrodesmus, anothergreen alga, becamequite abundantand apparentlymaintainedthe gas in abundance balanceduringthe fluctuations of Chlorella. The protozoan popula-

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tion generallyincreasedthroughout the experiment. Small snails appeared in the feces-fedsystemand were eatenavidlyby the fish. Sincethe weightof the fishsystemwouldhaveto be minimized, the relationships betweendensityand growthwerestudiedin volumesrangingfrom3.5 to 17 liters. Higherdensities,44 g/liter, and largerfish, 18 g averageweight,wereobtainedin the 17-litercontainers. Although Tilapiacan growmuchlargerin ponds,they are readilystunted; the largerof the test fishweremuchlike sardines. Thesecontainers wereopen to the air and further aeratedwith compressed air througha smallfilter with activatedcharcoal,sand,and glass wool. Brineshrimpwerefed ad lib daily. The waterwas changedif it becamefouled,usuallynot more than once per day. Someof the fishweresteamedand eatenby the research team:they werebonybut otherwisehighly palatable. The experiments wereinterpreted as suggesting that wasteproductsand excess algaecouldbe usedto producean alternate animalfood for the astronauts without an excessive02 demandor weight load. It was reportedthat an unidentified astronautopinedthat fishingmightproviderecreational value,but that eatingfecesrearedfish wouldbe an unlikelypossibility. Algal-Mammal-Waste Cycling An algal-microbial-mammal system,termedthe "microterella," was morecoupledthan formersystems;the wasteproductsof the mice providedthe algal nutrients, and condensedwaterfromthe systemprovided waterfor the mice(33). The totalsystemwas sealedin a 38.5 literchromatography jar. Spacewas providedfor one to four mice to resideon a mesh floorabovethe algal-bacterial culture.This areawas equipped with facilitiesfor eating,drinking, sleeping,and recreation,as well as an air conditioningunit whose condensate provided water.Freshfood in the formof pastewas supplied drinking fromoutside the system;to somebatchesalgalpastewasadded.The algal-microbial community occupied5-7 litersandconsisted of Chlorellapyrenoidosa (90-95%), Chlorococcum sp. (4-9%), Scenedesmus obliquus bacteria. (1%),and fecal-derived The objectives in these studieswere(a) to achieveconditions for maintaining a constant02/C02 ratio, (b) to test for limits of duration,and (c) to minimizethe supportsystem necessaryto maintainthe balance. The firstphase,reported in moredetail in Oswaldet al (70), providedinsights into the limits of regenerative systems.The initialseriesof experiments all led" to rapidincreases in CO2anddecreases in 02. Variousmodifications in retention time of the continuous algal-microbial to speedgrowthrateand 02 per cell community hadlimitedeffect.Reduction of thedepth(andvolume)of the culturewasbeneficial since in the shallowerculturea greaterportionof the algal populationwas well illuminated. theaddition Ultimately, of morelights,especially fromthe top,enabled a 5 literalgal-microbial to support community the 02 requirements of a mouseand its waste products. The secondphasewas concerned with the durationof gaseousregulation by the use of mouse wastesas the majornutritionfor the algal community. The culture fluidremovedin the continuous operationof the algal microbialcommunitywas andthe liquidreturned centrifuged to the unit.The unitfunctioned wellfor 10days;

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within 4 days after sealing the 02 rose to 25%, during the succeeding 5 days it varied from 23 to 25%, and on the 10th day it began to drop. At this time the culture had begun to turn yellowish green. Magnesium and phosphate were added, and within 2 days the culture became deep green and the 02, which had been 19% on the 11th day, was up to 25%. During the remainder of the 6 week run magnesium and phosphate were added routinely each 5th day, and the 02 remained between 23 and 25%. The run was terminated at the end of 6 weeks because Golueke & Oswald (33) felt that no further information would be gained. Additional experiments resulted in reduction of the volume of algal-microbial community required (33). Two points deserve examination: (a) a small volume was more effective than a large one, and (b) the microbial uses of 02 probably balanced the ultimate composition of the gas around 25% 02. A large volume of culture (143 ml/g mouse) was offset by inadequate light penetration, so that a smaller volume, 25 ml/g of mouse, was as effective. A large part of the increase in yield was reported to be due to an increase in the portion of the culture receiving an optimum amount of light. The potential self-regulationof the 02 in the mid-20% range is in interesting contrast to the 63% 02 of Bowman & Thomae (9). The further suggestion of homeostatic properties of such systems was made by Golueke & Oswald (33): The considerable influenceof carbondioxide on the system was demonstrated in experiments elsewhere reported et al, 1962)concerning (Oswald transfer functions with relationto step inputsof carbondioxide.The step inputswere accomplished by first removingone of two mice from the microterella when the system was in a state of withrespect equilibrium to gas;andthenrestoring the mouseafterthe oxygenconcentration hadagainreached Theimmediate equilibrium. in bothsteps. wasevaluated response Directlyafterremoval of the mouse,the oxygenconcentration dropped. However, after an interval of an houror so, the oxygenconcentration againreached equilibrium, usually at a level close to that beforeremovalof the mouse.Restoration of the mouse again abouta declinein oxygenanda repetition brought of the pattern observed uponremoval of the mouse. Apparently, the algalculturebuildsup an algalpopulation proportional to the carbon dioxideoutputof the micepresent in the unit.Whenthe mouseloadis suddenly reduced, the balancebetween carbon dioxideandalgalpopulation is upset,algalactivitydeclines, and oxygen production drops.Gradually, however,a new equilibrium is reached.The suddenrestoration of the originalmouseload, and consequent augmentation of carbon dioxideinputagainupsetsthe equilibrium. But, as before,the cultureadjustsitselfand the originalequilibrium is resumed. The abilityof the systemto adjustitself shows its homeostatic properties. (33, p. 526) Unfortunately, I have not been able to locate the experiments described above in my copy of the report cited. Perhaps that experiment was described in another of the progress reports from that laboratory. The nutritional closure of the system, though incomplete because the mouse was not fed entirely on algae, suggests that nitrogen as well as magnesium and phosphate might become limiting. The deficiency of nitrogen in their budget was due largely to the growth of young mice so that less nitrogen was returned to the algal culture than was being consumed (70).

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The authors made special mention of the pleasant woodsy odor of the microterella. Implications Crudeas these studiesof coupledsystemswere, they have several implications for a theoryof closed ecologicalsystems. 1. Continuous in which light is not incompatible with algalculture; experiments did not indicateany 24 hr periodicity withreduced gas wasmonitored continuously as mightbe expectedif internalmetabolicclocks regunighttimephotosynthesis, latedphotosynthesis or if metabolites produced duringthe darkwerenecessary for the next day's photosynthesis. Variationsin growth did occur in these coupled systems,as they did in Krauss'Recyclostat(41), but none of the authorsreported diurnalcycles. 2. The maintenance of the 02 concentration in the lower20% rangeis probably not due to any universal homeostatic mechanism of algal culturesalone,but may be a functionof algal-sewage communities, provided that light is adequate. Sucha mechanism mightbe hypothesized since high 02 concentration is knownto inhibit photosynthesis. Photosynthesis hadbeenreported to decrease whenthe culturewas in equilibrium with 02 pressures above 160 mm Hg (91). WhenCO2is in limited supplyits production by mice or microbesmight controlphotosynthesis. 3. Carbon did notappear monoxide to be a problem exceptin theblue-green algal culturesystems. 4. Self-inhibiting did not limit the recyclingof usedalgal culture algalproducts fromresultsobtained liquid,as had beenpostulated by Pratt(73) froman apparent autoinhibitor he called "chlorellin." Burlew(12) suggestedthat chlorellin,if produced,might be destroyedby the bacteriain contaminated cultures,and that an betweenthe algal and bacterialgrowths. ecologicalbalancemight be established Krauss(42) failed to find any evidenceof an autoinhibitor in his Recyclostat,in whichbacteria-free The autoinhibitor has not been algalculturesweremaintained. in Chlorella demonstrated to Myers,in any algalstrain pyrenoidosa, nor,according other than that used by Pratt(53). While closed and publishedby engineersand plant ecologicalsystems were being researched in the late 1950sandearly1960s,at leasta few ecologists triedto make physiologists someinput.Particularly H. Odum,his studentR. Beyers,andlaterE. Odumargued thatnaturally derived becauseof their systemswouldbe morelikelyto be successful information contentand highlyevolvedregulatory mechanisms. To increasecommunication betweenthe engineers and facilitateunderstanding and ecologistsand betweenboth groupsand the generalacademic biologicalcoma symposium, at the 1962 munity,I organized "Space Biology:Ecological. Aspects," Instituteof BiologicalSciences(AIBS) at OregonState meetingof the Aimerican Corvallis. The symposium University, was highlysponsored by variousAIBS organizations-the EcologicalSociety of America, AmericanSociety of Zoologists, BotanicalSocietyof America,Phycological Societyof America,Societyfor Industrial Microbiology, and AmericanFisheriesSociety,but not by NASA-and was well attended.
THE HOLISTIC APPROACH: THE ECOLOGISTS' CONTRIBUTION

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As far as I can judge, the symposiumand the subsequentpublicationof the Someecologoal;opinionsremained unchanged. presentations (79) achieved neither gists expressedskepticismthat algae could grow and survive at the high light to Matthern & Koch) (47) or under intensities reported (52,000lumensaccording continuouslight. Most of the engineersperceivedthe ecologistsas introducing unfeasible demands,such as a need for 2 unnecessary complexityand presenting of acresper man (61) or the need to initiatestudiesto gain furtherunderstanding naturalsystems(71). at the symposiumhas servedas a record. The publication of the presentations J. Myers(54) openedthe meeting andstatedthe differing pointsof viewof ecologists I compared and engineers. the presentsubsystem planswith the trophicorganizabalanced tion and presented the research described earlier(81). Beyers(5) reviewed and that the patternsof net photosynthesis aquaticmicrocosmsand emphasized or weresimilarregardless of majororganisms, temperature, nighttimerespiration the spacevehiclelife-support systemas a specialapplisalinity.H. Odumanalyzed he statedthat the cationof the moregeneralecosystems theory(61). In particular, of lightutilization dictatesthathighefficiency efficiency-power reciprocity principle cannotbe achievedwith maximalrate of output.He arguedthat the hiddencost of maintaining artificial algal systemsis reallygreaterthan the total cost of using of a system a multispecies, ecosystem.He proposedconstruction self-regulating from multipleseedingfromnaturalsources,which wouldbe unstablefor about2 andhighlymetabolic. Tischer self-sustaining, weeks,but wouldthenbe competitive, & Tischer(89) presented on the productionof Daphria, fungi, and experiments fromurineandfeces(reviewed bacteria earlierin this paper).Patten(71) presented tool for understanding naturalecosystemsand information theoryas a necessary & Koch (47) described the spacesystems.Matthern probably for wiselydesigning the uses of algaeand of massalgalcultures,and Wardet al presented engineering that of the other plants in space systems(96). Of the engineers'contributions, "Microterella" of Golueke & Oswaldwas most closely relatedin concept to a hereas a coupled ecosystem[thoughit wasreviewed naturally derived,multispecies includedan engineering approachto ecological subsystem(33)]. The symposium considerations of a permanent lunarbase(38) as well as a planfor the investigation was cited by life. Much of the presentation at this symposium of extra-terrestrial Cooke,Beyers& Odum(19) and by Cooke(18). The only contributions likely that I wasableto findin anyliterature by ecologists in the spacefieldwere(a) by E. Odumat the Discussion to be readby researchers on Regenerative in 1963,heldwithina Conference on HumanEcology(10), Systems Systems (b) a presentation by Cooke,Beyers& E. Odumat the Bioregenerative "Foods in 1966(19),andpossibly Conference at the symposium (c) my presentation in Space"(82), which marginally hinted at some ecologicalconsiderations. At the Regenerative SystemsConference,Brown (10), the discussionleader, indicatedthe lack of input from ecologistsand attributedit to the opinionthat the apecologistsdid not considerit part of their bailiwick.E. Odum presented folproachof H. Odum& Beyersof multipleseedingfrom naturalcommunities Brown lowed by closureand self-regulation, but it was met withoutenthusiasm. to space opinedthat such studiesare of interestto the ecologistbut not applicable

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systems.Still later, E. Odumreiterated the advisability of using stabilitymechanismsthat had evolvedin diversenaturalsystems,and Brownagainrejectedthis approach. FremontSmithsupported the ecologists'effortsto the extentof advocatingpursuit of bothapproaches, although no practical use wasyet apparent, "because the resultsthat may come of this line of thoughtare unpredictable . . ." Cookepresented the paperby Cooke,Beyers& Odumentitled"TheCasefor the Multispecies Ecological SystemwithSpecialReference to Succession andStability," at the Bioregenerative SystemsConference sponsored by NASA in 1966(19). In it the authorsstated that the engineering approachof testing and later assembling separate subsystems wouldsurelyfail, whereasthe ecologicalapproach of allowing groupsof speciesknownto occurtogetherto reassemble and reorganize in a new environment into an integrated, self-maintaining system,an ecosystem,wouldprovide the greatestopportunity for developinga successfullong-termlife-support system.Theysuggested the multispecies climaxecosystem as proposed by H. Odum (61). Theystressed thatlow efficiency withhighstabilitywouldbe feasible, whereas high-efficiency monocultures wouldrequire excessive external controlrequirements. Theydrewheavilyon the food webstudiesof Paine,the microecosystem studiesof Cookeand Beyers,and the trendstudiesof stagesof ecosystemsby E. Odum. Thediscussion afterthe meeting reflected the antagonism amongthe otherparticipants.The followingobjections werestated:this was merelyrewording the process of one-by-one selectionof components; a self-regulating ecosystem wouldnot necessarilyensurethe survival of anyone component, e.g. man;the spacesystemis being supported by earthtechnology and it does not haveto includeall of the functional components of the earth;monoculture is the basis of most of our agriculture and necessaryfor even a lunaror Martiancolony;the inclusionof 20 specieswould require210 experiments as a minimum; and it was questionedwhetherthe mixed culturesof Oswald& Goluekeweremorestablethan the two-component mousealga systemsof Myersor the single-algaculturesof Krauss.In spite of Cooke's rebuttals, the participants remained opposedto the conceptof a systemdeveloped by naturalseedingand self-controlled successionto an operational climaxstage. E. Odum,Beyers,andCookedid havea NASA research contract,NsG 706/11003, but it was terminated prematurely. This was the only work on ecological ecosystemssupported to my knowledge.[I had a small grant(NsG 519) to study the feasibility of usingthedevelopment of Daphniaundernull gravity as a potential Biosatelliteexperiment,but the experimentwas judged too complex because I plannedto use a lightedcultureof algae as a sourceof food and 02.1 ChemosyntheticRegenerative Systems With the increasingdisenchantment on the part of NASA with photosynthetic systemsbecauseof theirlow efficiency of electrical the Hydrogenomonas utilization, bacteria systembeganto look morefeasible.Electrolysis was deemedmoreacceptablethan photosynthesis becauseof its greaterefficiency Of02 production per unit of electricity. Bacteria of thegenusHydrogenomonas arecapable of utilizing gaseous hydrogen as an energysource,CO2as a carbonsource,andureaor ammonium salts as a nitrogensource.Tracematerials must also be suppliedin a growthmedium.

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cells The CO2and urinecouldbe recycledif H2weresuppliedto producebacterial the watercould and water:6H2+ 202 + CO2 (CH20) + 5H20. Subsequently be electrolyzedto supply 02 for the astronautsand to resupplyH2 for Hyculture.The cell mass might be a potentialfood. Obviouslythere drogenomonas but, as in the case of wouldbe a potentialsavingin electricalpowerrequirements; the algal culture,could a biologicalsystemhave adequatereliability? systemssparkeda numberof studieson NASA's interestin Hydrogenomonas Systems(56) in Bioregenerative growthand food value.Manyof thesearereviewed of Drakeet al (25), the papers System(55),especially LifeSupport andin TheClosed in 1972of (29) in the latter.Publication Jogowet al (36), and Foster& Litchfield Culture" eutropain Submerged of Hydrogenomonas the paper"MassCultivation by Litchfield(44) suggeststhat NASA is still pursuingthis approach.He cites animaltrials.A coupledculturehumanfeedingtrials,but successful unsuccessful no closedsystems not to havebeenstudiedyet;therefore electrolysis systemappears have been tested. relatedto spaceapplications The aboveis a roughsketchof the U.S. literature literaRussian the verysubstantial ignoring completely systems, of closedecological can be ture that appearsto follow the same trends.Access to the total literature key words. For a searchfrom NASA using appropriate obtainedby requesting and NASA indexed358 references, instance,usingthe key word"Bioregeneration" literature 461 more(58).TheRussian it indexed Systems" Ecological under"Closed is especiallywell coveredthere.The NationalTechnicalInformation (translated) Ecological on the title "Closed hundred references several (92) alsosupplied Service ProtectionAgency for is tenderedto the Environmental Systems"(appreciation supplyingthis material). NON-SPACE-RELATED STUDIES communithe studyof natural by the spaceprogram, In spiteof the lackof stimulus are not Whilenaturalsystemns ties has movedtowarda systemsanalysisapproach. life-supregenerative neitherwere the photosynthetic closed thermodynamically, Both are concernedwith the processeswithin an interacting port experiments. system so definedthat inputs and outputs can be measured.Two experimental to the study of naturalcommunitiesare especiallyrelatedto closed approaches ecologicalsystems:metabolismof naturalcommunitiesand microcosmstudies. of eithertopic. Spacepermitsonly a cursorytreatment Metabolism of Natural Communities e w'l tecognized as study (%W) the SWiey Spfimgs The zornireefstudy (63) nrld Numerousotherstudieshave used metabolism. classicsin the studyof community ratios apthe same approach(20-23, 64-66). P/R (photosynthesis/respiration) and lowervalues,i.e. proportionally maturecommunities, proach1 in undisturbed, seem mechanisms are foundunderorganicinputs.Homeostatic more respiration, to exist. It would be interestingto examine the data in sufficientdetail, or to new studies,to see whetherthe P/R ratiowouldbe greaterthan 1 with undertake

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the removal of organic material, e.g. fishing, and with the replacement of equivalent inorganic nutrients. Community metabolism deserves a review of its own. More detailed studies of community metabolism, or more particularly carbon flow, have been published by Saunders (77) and have been presented by Hobby and also by Wetzel (personal communications). These three authors have been comparing their results to determine whether different kinds of lakes have the same general patterns. Another approach to community metabolism was made by Olson (69), who used system analysis to examine the balance between producers and decomposers and the subsequent storage. System analysis is being used extensively now.

Microcosm Research
The research on microcosms has been reviewed recently by Cooke (17); for reasons of space, the studies in that review will not be discussed here. After reviewing the information on diurnal metabolism, ecological succession, P/R ratios, phosphorus cycling, and maturity and stability, Cooke concluded that these laboratory microcosms have demonstrated certain regulatory properties of ecosystems. Because microcosm studies require large populations of numerous interacting organisms, most have dealt with microbial communities. The entire symposium

publication Structure andFunction of Freshwater Microbial Communities (13) may

serve as a review itself. In that volume I presented the results of a continuous gnotobiotic ecosystem consisting of an alga, a herbivorous protozoan, and two bacteria (83). Since then I have simplified the continuous cultures to only two organisms, the alga and the herbivorous protozoan. The relationship between the nutrient, alga, and protozoan has been modelled mathematically with the help of D. McKenzie of our laboratory (84, 86). I am now in the process, with N. Pearson, of attempting to verify the predictions of the model. In the course of these studies I have become aware of another experimental-mathematical treatment of stability relationships of continuous predator-preycultures. These studies (14, 15, 24, 26, 39, 80, 90) are heterotrophic, being based on an organic compound and bacteriaprotozoa interactions. In contrast, my system starts almost devoid of organics and the protozoa depend on the photosynthesis and growth of the algae as their sole source of food. I believe a review on predator-prey continuous cultures is in preparation.

Closed Microcosms and Balanced Aquaria Studies3

Most of this work, including my own, appears to be unpublished. Both Beyers and Cooke have made observations on closed systems, but have not completed studies 3Since the review,two papers on massandenergyflowin closedecosystems have preparing beenfound.Ulanowicz(100)proposed a mathematical for linearsystemsrelating framework theparameters of population to thoseof energy more dynamics flow;May(101)hasconsidered for ecosystem complexwebs.They statedthata necessary andsufficient condition stabilityis thatenergy contentperunitbiomass increase in a hierarchical fashion as oneclimbsthetrophic ladder. Otherinsights on trophic or observational relationships areindicated. No experimental resultsare considered. Thesemodelsmightprovidethe basisfor validation experiments.

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systems suitablefor publication.E. Odum (59) picturedseveral demonstration for teachingpurposes,but no data were shown.Two comestablished apparently involved1. a seriesof pairedalgaland bacterial pletedstudiesfrommy laboratory derivedculturesenriched culturesconnectedonly by a gas bridge,and 2. naturally with varyingamountsof nutrients. I was unableto findany detailedresearches. aquaria, On the subjectof balanced of a 12 gallon E. Odum (59) cited a reportby Waringtonof the establishment to be impossible, aquaria statedbalanced system.Atz (3) had apparently aquarium them(citedin 62).Odum&Johnson havetriedto establish at leastas mostaquarists but that fishareimpossible, (62) agreewith Atz that smallsystemswith numerous is not impossiblein concept,given an adequatebiomassof the sealed aquarium forsalein 1973by two wereprepared aquaria anddecomposers. Balanced producers state, AnnetteNormanand SarahWilson.Most of their women in Washington systemswere gallonjugs with sand, water(2/3), air (1/3) and severalplants,pond did not persist,as they were detritussnails, and a pair of guppies.Zooplankton The rapidlyeaten.The fish tendedto live about4 monthsbut rarelyreproduced. snails survivedfor about 6 months in the system they gave me, and the plants healthyafter 10 months.Their resultsappearto be typicalof most atappeared tempts. SUMMARY theholisticapproach to combine nevermanaged of closedecological systems Studies of the few ecologists active in that field with the subsystemsapproachof the studies offered The only hypotheses by the engineering and physiologists. engineers for the controlof 02 mightbe responsible mechanism were(a) that a homeostatic in the low 20% range,and (b) that some of the supportfor a higher concentration to growfungior fish.The trophiclevel (man)mightcomefromusingfecalmaterial forests,and desertsare now receivingincreased detritalcycles of lakes,estuaries, and I have P. Bissonette BiologicalProgram. study,especiallyin the International been exploringthe hypothesisthat much detritalmaterialin lakes becomesconeatenby fish. aresubsequently andthe chironomids biomass vertedinto chironomid of agriculture mayyet be usedin the production The massalgalculturetechniques of the production the smallcells fromthe liquidis a problem, or, since harvesting (85). aquaticfood for aquaticfilterfeeders systemsof thusfar withoutbiologicalregenerative The spaceefforthas managed or chemosynthetic type. Sky Lab, with its crew of three eitherthe photosynthetic matewithstoredexpendable managed in excessof 2 months,apparently remaining waterfrom the fuel cells (28). rials and probably inherent of the self-regulation Sealedecologicalsystemsdo permitexamination Gaseousexchange witha finiteset of chemicalresources. in a givenset of organisms to measure directly,and estimatesof 02 and CO2transfer is difficult and expensive of concentraor estimations measurements from02 diffusion andpH changerequire Nitrogentransferis even more difficultto tion gradientsand mixingcoefficients. techniques N2 fixationor release.Respirometry unlessthereis significant measure

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introduceartifactsbecausethey requiredense concentrations of organismsand extremepH buffering. The inclusionof an atmosphere in a sealedsystempermits the water-atmosphere diffusion and storagefunctionsthat occurin the biosphere. In a sealed system, transientconcentrations Of 02 and CO2 can be monitored continuously and can providea more detailedrecordof communitymetabolism. This approachmay yet providean excellenttool for studyingtotal community responsesto manipulations.
Literature Cited 1. Adams,C. C. 1963.Nutritional aspects of space flight.Medicaland Biological Problemsof Space Flight, ed. G. H. Bourne,Chap. 10, 237-44. New York: Academic.289 pp. 2. AeronauticalSystems Division, Dir/ Aeromechanics, Flight Accessories Lab., Wright-Patterson AFB, Ohio. 1963. Atmospheric Control Systems for Space Vehicles.Rep. ASD-TDR-62527, Part I. Arlington, Va.: Armed Serv.Tech. Inf. Agency.247 pp. 3. Atz, J. W. 1949. The balanced aquariummyth. Aquarist 14:159-60, 179-82 F. B., Taufman, 4. Benjamin, W. I., Helvey, W. M. 1962.Biologistics for Space Systems Symposium,242-64. Aerospace Med. Div., 6570th Aerospace Med. Res. Lab., Wright-Patterson AFB, Ohio AMRL-TDR-62-116 5. Beyers, R. J. 1963. Balancedaquatic microcosms-their implications for space travel. Am. Biol. Teach. 25: 422-29 6. Bieberdorf, F. W. 1962.A Studyof MicrobiologicalWaste TreatmentTechniques.USAF 6570thAerospace Med. Res. Lab., Wright-Patterson AF Base, Ohio, AMRL-TDR-62-142.83 pp. 7. Bovee, H. H. et al 1962. Largealgal systems.See Ref. 4, 8-19 8. Bowman,R. O., Thomae,F. W. 1960. An algaelife support system.Aerospace 82 Eng. 19(12):26-29, 9. Bowman,R. O., Thomae,F. W. 1961. nontoxicsupportof animal Long-term life with algae. Science134:55-56 10. Brown,A. H. 1966. Regenerative systems. Proc. Ist Int. Interdisciplinary Conf:,Princeton, NJ., October 13-16, 82-119.New 1963,ed. D. H. Calloway, York:New York Acad. Sci. 11. Burlew,J. S. 1961.Algal Culture from to Pilot Plant. Washington Laboratory DC: CarnegieInst. Publ. 600. 357 pp. (Firstprinting1953) 12. Burlew, J. S. 1961.Current Statusof the LargeScale Culture of Algae. See Ref. 11, 3-23 13. Cairns,J. Jr., Ed. 1969. TheStructure and Function of FreshWater Microbial Communities. Res. Div. Monogr. 3, Virginia Polytech.Inst.andStateUniv., Blacksburg. 301 pp. 14. Canale, R. P. 1970.An analysis of models describing predator-prey interaction. Biotechnol. Bioeng.12:353-78 15. Canale,R. P. 1973. Experimental and mathematical modeling studies of protozoan predation on bacteria. Biotechnol. Bioeng. 15:707-28 16. Conrad,H. M., Johnson,S. P. 1965. Problems associated withthe utilization of algae for food. Activ. Rep. 17(1): 79-85 17. Cooke,G. D. 1969.Aquaticlaboratory microsystemsand communities.See Ref. 13, 47-85 18. Cooke, G. D. 1971. Ecologyof space travel. Fundamentals of Ecology, ed. E. P. Odum,Chap.20, 498-509. Philadelphia:Saunder 19. Cooke, G. D., Beyers, R. J., Odum, E. P. 1968.Thecaseforthemultispecies ecological system, with special reference to successionand stability.Conf: Bioregenerative Systems, Washington, D.C, November 15-16, 1966, 129-39. NASA SP-165 B. J. 1965.Evidence 20. Copeland, for regulation of communitymetabolismin a marine ecosystem. Ecology 46(4): 563-64 21. Copeland, B. J.,Jones,R. S. 1965.Comin somehypersaline munitymetabolism waters. Tex.J. Sci. 17(2):188-205 22. Copeland,B. J., Whitworth,W. R. 1963.Oxygenmetabolism of fourOklahomafarmponds.Inst. Mar.Sci. Tex. 9:156-61 23. Copeland,B. J., Davis, T. C. 1964. in ecosystems Community metabolism oil refinery effluents. Limnol. receiving Oceanogr. 9(3):431-47 24. Curds, C. R. 1971. A computersimulation study of predator-prey

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relationships in a single-stage continuous-culture system. Water Res. 5:793812 25. Drake, G. L., King, C. D., Johnson, W. A., Zuraw, E. A. 1967. Study of life-support systems for space missions exceeding one year in duration. The

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26. 27. NASA SP-134 Drake, J. F., Jost, J. L., Fredrickson, A. G., Tsuchiya, H. M. 1968. The food chain. See Ref. 19, 87-95 Eley, J. H. Jr., Myers, J. 1964. Study of photosynthetic gas exchange, a quantitative repetition of the Priestley Experiment. Tex. J. Sci. 26:296-333 Fink, D. E. 1973. Flexibility stressed in Skylab hardware. Aviat. Week Space Tech. 98(15):52-55 Foster, J. F., Litchfield, J. H. 1967. The effects of controlled environment on the growth of Hydrogenomonas bacteria in continuous cultures. See Ref. 25, 201-12 Gafford, R. D., Bitter, H. L., Adams,

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