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W0I. C. cCh. NO!.2J, PO.^, pp. JV~b2,1VV2.

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PROCESSES AND MODELLING OF
NITRIFICATION DENITRFICATION
BIOLOGICAL EXCESS PHOSPHORUS
REMOVAL SYSTEMS - A REVIEW
M. C. Wentzel, G. A. Ekama and G. v. R. Marais
2!~1222 1JUU
Copyright G VV2PWKL
Department of Civil Enlineerinl. University of Cape Town, ROIt/I'/J(lSciJ, 7700, SOl/II!
Africa
ABSTRACT
This paper reviews developments in modelling the kinetics of activated sludge systems: Completely
aerobic nitrifcation, anoxic/aerobic nitrifcation denitrifcation ( ND), and anaerobic/anoxc/aerobic
ntrifcation denitrifcation biological excess phosphorus removal ( NDBEPR) systems. The paper
hghlights the progress in developing a general NDBEPR activated sludge kinetic model - development of
polyP organism enhanced cultures, their kinetics, simplifcation of the kinetics for enhanced cultures under
constant fow and load conditions, extension of the Simplifed model to mixed culture NDBEPR systems
under constant fow and load conditions, integration of the polyP organism enhanced culture kinetics with
the ND kinetics to give a general NDBEPR kinetic model for cyclic fow and load which incorporates the
increased specifc denitrifcation rates observed in NDBEPR systems compared to ND systems. Areas of
research that require attention to complete the development of the general NDBEPR knetic model are
identifed - denitrifcation by polyP organisms, caibration and verifcation of the model for cyclic fow
and load, etc.
KEYORDS
Kinetics, steady state models, biological excess phosphorus removal, nitrifcation, dentrifcation, nutrient
removal, enhanced cultures, mixed cultures, readily biodegradable COD, slowly biodegradable COD.
INTRODUCTION
Since the early 1970s signifcant developments have taken place in the activated sludge method of
treating wastewaters. The function of the single sludge system has expanded from carbonaceous energy
removal to include progressively nitrifcation, denitrifcation and phosphorus removal, al of these
mediated biologically. These extensions have been accommodated through manipulation of the system
confguration - incorporation of multiple in-series reactors, some aerated and others not, with various
inter-reactor recycles. Not only have the system confguration and its operation increased in complexity,
but concomitantly the number of biologica processes infuencing the efuent quality and the number of
cmpounds involved in these processes, have increased. With such complexity, it is no longer possible to
make a reliable quantitative, or sometimes even qualitative prediction as to the efuent quality to be
epected from a design, or to assess the effect of a system or operati6nal modifcation, without some
model that simulates the system behaviour accurately. This paper reviews progress in the development of
such a model for the single sludge nitrifcation denitrifcation biological excess phosphorus remova
(NDBEPR) activated sludge system.
EARY DEVLOPMENTS
In the wastewater treatment feld, in response to the pressure for solutions to immediate problems,
ful-scale application and implementation of a treatment system often precedes the science and
understandng of how the system works. This certainly has been the case for the NDBEPR activated
sludge system.
59
60 M. C. WENTEL 0I al.
The NDBPR activated sludge system was develcped in II4 by Barnard, thrcugh the inccrpcraticn of
biclcgica| excess phcsphcrus remcv [BPR) in the nitricaticn deniiricaticn [ ND) activated sludge
system. Initial ccnceptualitaticn cfNDBPR syste behavicur extended li tt|e beycnd reccgniticn cf [ I )
the necessity cfan anaercbic]aercbic sequence in the syste, and (2) ihe adverse in0uence cfnitraie
recycled tctheanaercbictcne, Barnard [II4) andlaterccnrmedby Barnard II I I) and Nichclls
[ II). The reascns fcr these factcrs were nct understccd, but experienta| cservaticns indicated that
the presence cf an anaercbic tcne stiulated BPR, and that high niirate discharged tc the anaercbic
tcne signicantly reduced BPR. The NDBPR system ccnguraticn that was develcped was the
-stage Hcdied Bardenphc. Ths-stage system was develcped frcm the 4tageBardenphc ND system
[Barnard, II4) which ccnsisted cf ineries primary ancxc, aercbic, seccndary ancxc and re-aeraticn
reactcrs, with a recyclefrcm the aercbic tc the primary ancxc reactcr and an under0cw recycle frcm the
settling tanktc the primary ancxc reactcr, see Iig Ia. The cb]ective cfthe4t age system was tcmtrify
the ammcniain theaercbic reactcrs and tc denitrify theresultant nitrate in theancxic reactcrs, tc givea
mnmal ammcniaandnitrateccncentraticn in the eMuent . In the tage system an "anaercbic" reactor
was inccrpcrated ineries upstreamcftheprimary ancxc reactcr, withihe under0cw recycle and in0uent
dscharging tc the anaercbic reactcr, see Iig Ib, in this system it was expected that the nitrate in the
eMuent , and hencein theunder0ow recycle, wculd be sc lcw that its e!!ect cn reducing BPR wculd be
salI. Design procedures devclved toempiricly based estimates |ct sitiag denitricaticn and anaercbic
reactors interms ofnominal hydraulic retenticn timc. Nc raticnaI ethcd fcr premcting 00D, nitrcgen
and phosphorus removal was avlable, in design, remcvals were estimated l argely frcmexperiencegained
inoperating experimentalsystems similartothe prcpcsed systes .
The lack of a method for predicting removs focused attenticn cn deveIcping models to identify and
describe the kinetic behaviour of the different processes acting in the NDBPR system, 00D remov,
mtrication, denitrication and BPR. The kinetic descripticn c| these prccesses evolved in an
apprcximately sequential manner, in aercbic, ancxic]aercbic [ND) and anaercbic]ancxc]aercbic
[NDBPR) systems. Thus, it i snot possible toreview deve|cpments ia describing the kinetic behavicur
ofthe NDPRsystemwithcut dealingwi ththeantecedent systems
AEROBIC NITRFICATION SYSTEM
Steady state aeobic model
!n I16 Haras and Bkama proposed a steady state model for the aercbic nitrication system under
constant 0ow andloadcondticns. This steady statemodel constituted a develcpment frcm a number of
previcusmcdelsfcr carbonaceous and nitrogeno's matericcnversicn andremcv.
Caronaceou materal Hcdelling cf carbcnacecus aterial ccnvetsicn ccnfcrmed principally tc the
propcsals cfHcKinney [ I 62) and HcKinney and 0cten [ I). These researchers proposed three mixed
liqucr vclatile solids fractions, active, endcgencus-inert and inert, the Iast named derived frcm the
in0uent . Iurther, they proposed [I) a nlaticnshp between the mass c|substrate utilited and the active
mass cf crganisms synthesited, and an expressicn fcr the rate cf crganism mass synthesis, (2) an
accumulaticncfvclatileendcgencus-inert sclids [endcgencus residue) asscciated withactivemasslcss due
to endcgencus respiraticn, and an expressicn fcr the rate cf active mass lcss, [) a relaticnship between
the cxygen demand and the crganism active mass synthesited, [4) a relaticnsip between the cxygen
demand and the active mass lcss due tcendcgencus respiraticn, and (5) an accumulaticncfinert vclatile
solids due tc t he presence cf tHs material in the in0uent. Harais and kama [ II) accepted these
proposs, with the exception of HcKinney's propcs fcr the rate cf synthesis cf active mass. Instead
they accepted Iawrence and Hc0arty's [ II0) prcpcs linking the specic crganism grcwth rate to the
concentrationofsubstratesurroundngtheorganismviatheHcncd relaticnship.
Haras and kama [ II6) re]ected the bicchemical cxygen demand [ B0D) as a suitable parameter fcr
dening t he carbcnaceous material. Instead they accepted t he e|ectrcn dcnating capacity cf the
carbonaceous material in its eqmvalent fcrm, thechemicalcxygen demand [ 00D). They prcpcsed that
thein0uent 00Dbedivided intothree fractions, [ I) bicdegradable, (2) unbicdegradableparticulate, and
[l) unbicdegradable soluble. The growth and endcgencus respiraticn prccesses were fcrmulated in terms
of the 00D, or, where apprcpriate, in terms cf an asscciated paraeter the vclatile suspended sclids
(VSS). Based cn these consideraticns Hars and kama [ II) deve|cped steady state equaticns fcr
single and in-seriescompletelymixedreactor activatedsludge systes teceiving aconstant 0cw and lcad,
i . e. equations for the active, endcgenous and inert vclatile scli ds ccncentrations and fcr oxygen
consumpticnrates due to synthesis andendogenous respiration, these as |uncticns ofsludge age [organism
retenticn time) . Development of these steady state equaticns was faoli tated by the observaticn that
understeadystateconditions thekineticswerevirtulycomplete andcculdbereplacedby stoichicmetric
rdationships, except for endogenous respiration the kinetics cf which were fcrUated according to the
proposalsofHcKinneyand0oten[ I) .
Nirogenou materal Harais and Bkama [ II) proposed that thc |auent nittcgen [N) be divided intc
NBEPR systms
61
fcur fracticns, I ) unbic1egra1able scluble, [ 2) unbic1egra1able particulate, [ l) bic1egra1able orgamc,
an1 [4) free an saline ammcnia. Again they 1evelcpe1 stea1y state equaticns fcr the utilitaticn cf
ammcnia fcr active cell mass synthesis , reease cf crganic nitrcgen 1ue tc en1cgencus respiraticn,
ccnversicn cf bic1egra1able crganic nitrcgen tc ammcnia, inccrpcraticn cf unbic1egra1able nitrcgen in
inert material, an1 ccnversicn cf ammcnia tc nitrate [nitricaticn) as functicns cf slu1ge age. The
ccnversicn cfammcnia tcnitrate require1 the inccrpcraticn cfa kinetic equaticn, fcr this purpcse they
cllcwed theHcnc1 apprcach, as setcut byDcwning, Painter an1Kncwles [ I4).
D_mcmodd
The stea1y state aercbic mc1el, when it was applie1 tc simulate single an1 ineries aercbic reactcr
systes un1er cyclic cw an1 lca1 ccn1iticns, gave pre1icticns that 1eviate1 signicantly frcm thcse
cbserved experimentally. The stea1y state aercbic mc1el ccul1 nct explain the cbservaticns ma1e in
experimental investigaticns by kama an1 Harais [ II), they fcun1 that aercbic ccmpletely mixe1
single reactcr systems at shcrt slu1ge ages [ I . -l. 1) uner 1ly cyclic square wave lca1ing ccn1iticns
[fee1 I2h cn, I2h cff) , exhibite1 aprecipitcus 1rcpinthecxygenutilitaticnrate [00R) cnterminaticncf
the fee1, see Pig 2. Tc explain this step change Dcl1 et al. [ I) prcpcse1 that the inuent
bic1egra1able C0D be 1ivi1e1 intc twc fracticns, rea1ily bic1egra1able C0D [ RBC0D) an1 slcwly
bic1egra1able C0D [5BC0D) - 1uring the fee1 peric1 the RBC0D is utilite1 at a very high rate an1
virtually ccmpletely, at fee1 terminaticn ad1iticn cf RBC0D alsc ceases causing a ccnccmitant
precipitcus 1rcpinthe 00R.
0cl1 et al. inccrpcrate1 RB00D an1 5BC0D in an aercbic nitricaticn activate1 slu1ge Hnetic mc1el
that gave very gcc1 pre1icticns cf the cbserve1 precipitcus 1rcp in 00R, an1 cf the 00D, 00R and
mtrcgen behavicur in varicus aercbic systems un1er cyclic 0cw and lca1 ccn1iticns. In the mc1el, they
pcstulate1 that the twc 00D substrate types, RB00D an1 5B00D, are acte1 cn in1epen1ently by the
tctal activecrganism mass inthe system.
Readily biodegradable substrate (RBCOD): The RB00D, they hypcthesize1, ccnsists cf sm simple
mclecules that can pass 1irectly thrcugh the cell wl [by passive cr active uptake) fcrsynthesis and
cxi1ative metabclism by the crgamsm. The rate cf synthesis cf active mass frcm the RB00D is
fcrmUate1 acccrdng tctheMcnc1 equaticnlinkingthespecicgrcwth ratecftheactivemasstcthe
RBC0Dccncentraticnintheliqui1phase. The reacticnrateisrapi1.
Slowly biodegradable substrate (SBCOD): The 5B00D, they hypcthesite1, ccnsistscflarger ccmplex
mclecules that cannct pass 1irectly thrcugh the cdl wall. 0tilitaticn cf this fracticn invclves fcur
1istinct phases; [ I ) enmeshment by the slu1ge mass, [2) a1scrpticn, l) extracellwar enzymatic
break1cwn cf the ccmple crganic mclecules tc simpler ccmpcnents whic pass directly thrch the
cell wall, an1 [1) metabclism cf the simpler ccmpcnents by the crganism. Pct phase the
enmeshment cfthe 5B00Dwas assume1 tcbeinstantanus an150 wasnctexplicitlymcdde. Pcr
phases [l) an1 [1) , the cverall reacticn rate is slcw an1 appears tc be limite1 by the rate cf
extracelluar break1cwn [hy1rclysis) cf the a1scrbe1 5BC0D materi [phase [l rather than the
rate cfmetabclism cf the hy1rclyze1 materi which passes thrcugh the cell wl [phase [1)] 5ince
hy1rclysis is the rate limiting prccess, abscrpticn an1 subsequent metabclism cf the y1rclysis
prc1ucts were nct exlicitly mc1elle1. Bence, the utilitaticn cf 5BC0D ccul1 be mc1elle1 as a
twc-stage prccess, I) a1scrpticn cf the material by the crganism ass, an1 (2) entymatic
break1cwn [hy1rclysis) cf the substrate. They hypcthesite1 that the hy1rclysis prc1ucts pass
1irectly tcthecrgamsm,nct tcthescluticn surrcun1ingthecrganism. -
Incnefurther aspect the 1ynamic mc1el cf Dcl1 et al. 1iffere1 substantively cm the exstin stea1y
state mc1els. In the earlier mc1els the en1cgencus respiraticn ccncept was use1 tc expn the
phencmencn cf active mass lcss, i. e. the re1ucticn in active vclatile mass with time. This prccess was
attribute1 tc an energy requirement fcr crganism maintenance, where a fracticn cf the crganism mass
dsappears tc prcvi1e this energy requirement . Determinaticn cf the rate cf mass lcss via cxygen
utilitaticn rate [ 00R)vs time plcts, measure1inbatch aercbic 1igesticn studes, shcwe1 that the speoc
mass lcss rate ccnstant was very high, .24 mgAV55]mgAV55{1, greatly exc1ing the specic rate
ccnstant chen cbserve1 fcr pure cultures, . 2. mgAV55]mgA V55]1 [HcKinney an1 0cten, I;
Hars an1 Bkama, I I) . Purthermcre, in a reactcr with sequential aercbic an1 ancxic states un1er
!This prcpcse1 mechanism fcr 5BC0D utilitaticn 1iffers cm that inccrpcrate1 in the IAWPRC Task
0rcup mc1el [Benze et aL, II). Inthe Task 0rcup mc1eitis prcpcse1 that the5BC0Dishy1rclyze1
extracellularly tc RB00D, which then is release1 tc scluticn, tc be utilite1 ccmpetitively by l the
crganisms via the mechanisms 1etle1 abcve fcr RB00D utilitaticn. Difculties with the Task Crcup
apprcach are enccuntere1inexten1ing the Task 0rcup mc1el tc inccrpcrate BPR. Preliminary results
frcm anexperimentalenqury intc the twc apprcaches appear tcfavcurthe 0CT apprcach [Massi11a et
aL, II). Theseccnsi1eraticnshavele1the00T 0rcup tcreverttcthecriginprcpcsscfDcld et at
[ I) .
Jhb1 b10-l
6
M. C. WL 0lal.
ccnstant cw an1 lca1 cr batch ccn1iticns, in situaticns where the ccncentraticn cf bcth cygen an1
nitrate were zerc fcr a peric1 [i e anaercbic state) , it was cbserve1 that when cxygen [cr nitrate) was
reintrc1uce1 the initial c

en [cr nitrate) 1eman1 increase1 signicantly, Warner et al. (986). The


increased cxygen [cr nitrate 1eman1 was fcun1 tc be equal tc the tctal 1eman1 that wc1 have been
expecte1 if cxygen [cr nitrate) had been available 1uring the anaercbic peric1. The en1cgencus
respiraticn apprcach ccul1 nci explain this cbservaticn Thus there was a nee1 tc1evelcp analternative
ccnceptualmc1el which ccul1 prcvi1e a reascnableexplanaticnfcrthehigh specic en1cgencus mass lcss
rate an1 reect the situaticn where aercbically [ancxically) generate1 crganisms were place1 in an
anaercbic statefcr ashcrt peric1 This le1 tc thefcrmulaticncfthe 1eath-regeneraticnhypcthesis.
Inthe 1eath regeneraticn hypcthesis anattempt is ma1e tc separate cut thereacticns which take place
1uring the crganism's "en1cgencus"phase Disappearance cflive active mass ishypcthesite1tcbe1uetc
thenet e0ect cf1eath [natural crpre1aticn) an1 regeneraticncfcrganisms. 0n 1eath thecell materi is
release1 thrcugh lysis, a acticn is unbic1egra1able an1 remains as an unbic1egra1able en1cgencus
resi1ue, the remaining fracticn isbic1egra1able an1 beccmes part cf the5BC0Din the liqui1, returning
tc the sae cycle cf adscrpticn, hy1rclysis an1, nly, synthesis cf new cel mass [ie. regeneraticn),
giving rise tc an asscciate1 synthesis cxygen 1eman1 The classic en1cgencus respiraticn cxygen
1eman1 ihus in effect is explaine1 as a resynthesis cxygen 1eman1. The main implicaticn cf this
apprcach is that "maintenance energy" per se [the cxygen requirement fcrmaintenance) isccnsi1ere1 tc
bescsmall that it can belumpe1with, an1ccmpletely swampe1 by, thecxygen1emanfcrthe synthesis
cfnewcellmassom the lyse1 substrate Pcrmulaticncfthisapprcachin1icate1 a specic 1eath ratecf
0.62 mgAV55mgAV55]1 giving with resynthesis a net specic active mass lcss rate cf 0. 24
mgAV5]mgA55]1 0sing either the classical en1cgencus respiraticn cr the 1eath regeneraticn
apprcach with aercLic systems gave virtually i1entical results, it was pcssible tcccmetca mcrepcsitive
1ecisicn as tcwhich apprcachissupericrcnlywhen 1enitricaticn an1 cr anaercbicstates wereinclu1e1
inthemc1el
ANOXIC/AEROBIC NITRFICATION DENITRFICATION SYSTEM
In XD systems un1er ccnstant cw an1 lca1 ccn1iticns , 5tern an1 Harais (1974) shcwe1 that
1enitricaticnin a plugcw primar ancxc reactcr tcck place in twc linear phases, a rapi1 rst phase
which persiste1fcra shcrt peric1 then terminate1, an1 a seccn1 slcw phasewhich ccntinue1fcrthe rest
cfthereienticn timeinthereactcr, in aplugcw secondar ancxicreactcrcnly cnelinear 1enitricaticn
phase was cperative, at a slcw rate abcut twc thir1s that cfthe slcw seccn1 rate in the pnmary ancxic
reactcr. kama et al. (1979) hypcthesize1 that the twc linear phases in the primary ancxc reactcr arcse
cm the utilizaticn cfthe twc bic1egra1able C0D acticns in the inuent, RBC0D an1 5BC0D; the
rst 1enitricaticn phasethey ccnnecte1 tc the RBC0D an1 the seccn1 tc 5BC0D. With regar1 tc the
slcw rate single 1enitricaticn phase cbserve1in the seccn1ary ancxic reactcr, they prcpcse1 this tc be
1ue tc en1cgencus mass lcss Base1 cn these cbservaticns they 1evelcpe1 a 1enitricaticn 1esign
prcce1urefcrin-series multi-reactcr systems un1er ccnstant cw an1lca1 ccn1iticns.
VanBaan1el et al. (1981) inccrpcrate11enitricaticninthesynthesis 1eathregeneraticnaercbicmc1ecf
Dcl1 et al., tc prc1uce a general nitricaticn]1enitricaticn [XD) activate1 slu1ge kinetic mc1e. V
Baan1el et al. fcun1 that the1enitricaticn kineticbehavicur ccuI1 be mc1ele1 in terms cfRBC0D1
5BC0D, an1 that the same fcrmulaticns prcpcse1 by Dcl1 et al. fcr RBC0D an1 5BC0D utilizaticn
un1eraercbicccn1iiicns ccU1 be use1 tcmc1el their utilitaticn un1erancxicccn1iticns, exceptthat the
rate cf5BC0D hy1rclysis]utilitaticn un1erancxicccn1iticnsn1e1 tc be re1uce1 tc abcut 1/3 cfthat
un1er aercbic ccn1iticns 0sing the gener nitricatic1enitricaticn mc1el, simlaticns cf the
1enitricaticnrespcnseinprimary an1 seccn1ary ancxic plu cwreactcrs pre1icte1near linear twcphase
1enitricaticn behavicur in the primary ancxic reactcr, an a single near linear phase in the seccn1ary
ancxc reactcr, as cbserve1 by 5ten an1 Harais (1974). Van Baan1el et al. (1981) ccndu1e1 that , inthe
plugcw primary ancxic reactcr the rst phase linear 1enitricaticn rate arcse cm utilitaticn cf
RBC0D]a1scrbe1 5BC0D an1 the seccn1 frcm utilizaticn cf a1scrbe1 5BC0D. In the pl;0cw
seccn1ary ancxic reactcr the single 1enitricaticn phase arcse cm utilitaticn cf a1scrbe1 B0D
generate1cmcrganism1eath Purthermcre,inapplicaticn tcsituaticns where ccncentraticnscfcxygen
an1 nitrate were zerc fcr a shcrt peric1 the mc1el was able tc simulate the cbserve1 increasein cxygen
[nitrate) 1eman1 when cxygen [cr nitrate) was reintrc1uce1 prcvi1e1 the 1eath regeneraticn apprcach
was use1 - thi s implie1 that the 1eath regeneraticn apprcach was supericr tc the classical en1cgencus
respiraticn apprcach, Warner et al. (1986).
Accepting linear 1enitricaticn phases, van Baan1el an1 Harais (1981), v Baan1el et a (1982) an1
kama et aL (1983) 1evelcpe1 a simplie1 stea1ystate mathematic mc1elfcrND systems. his stea1y
state mc1el prcvi1e1 gui1elines fcr the 1esign cf ND systems, fcr siting ancxic reactcrs, estimating the
1enitricaticnpctential, an1cthers.
NBEPR systms 63
k@caticncfND stmdystate mcdeltcNDBEPR svstes
In XBBPR systems which inccrpcrate anaercbic reactcrs, the simplie1 steady state 1enitricaticn
iheory was appIied unmcdied, ccmpariscn cf experiment an1 cculated 1enitricaticn respcnses in
steady state cperaticn appearedtc]ustlfy this applicaticn[vanBaan1elan1Mars,1981). The simplied
RB cdeI prcvlded estimates cf the denltricaticn pctentials cf ancxic reactcrs, this facilitated the
1evelopment cfXDBPRsystems tha: prcvided greater erbiIity an1 security in ensuring zero discharge
of nitrate to the anaerobic zone, the 0CT [Rabincwitt an1 Mars, 1980) and M0CT [ 5iebritt et aL,
1980) systems an1 later theJchannesburg system [Nichclls et al., 1987). he 1ifferent ccnguraticnsare
comparedinIig 1.
BEPR PARAMETER IDENTIFICATION
Baving deveIcped system ccnguraticns in which it was pcssibIe tc eIiminate the ccnfcunding eect cn
BPR cf nitrate recycled tc the anaercbic reactcr, it was pcssible tc i 1entify cther parameters that
imuence the magnitude cf BPR. Investigaticns using the 0CT type systems i1entie1 twc key
parameters that appeared tc inuenceBPR, the anaercbicslu1ge mass fracticn an1 the ccncetraticncf
hBCOD surrcunding the organisms in the anaercbic tcne With regar1 tc the fcrmer, increasing the
anaercbic mass fracticn led tc an increasein BPR, but at adecreasing rate. With regar1tc thelatter,
increasing the RBC0B ccncentraticn led tc an increase in BPR. In terms cf these parameters, the
inuence cfnitratecn BPRccuIdbedetermined, ifnitratei srecy
cle1 tcthe anaercbictcne,RBCOD is
utiIited preferentiIy with nitrate as electrcn acceptcr thereby causing a re1ucticn in RBCOD
concentraticnintheanaercbictcneandaccnccmitant reducticninBPRby thesystem. Base1 cnthese
concepts, retainlng the simplied XB thecry, design proce1ures were 1evecpe1 in which BEPR was
formulatedempiricaIIyintermscfthe twc key parameters an1 the masscfslu1ge[active, en1cgencus an1
inert) wasted perday [ 5iebritt et al., 1983; kama et ai. , 1983; WRC, 1984).
ssentiIy up tc this time, descripticn cf XDBPR system behavicur 1i1 nct reccgmte the presence cf
any specic crganism implicated in BPR. The 5iebritt et al. mc1e in fact ccnsi1ere1 the activesludge
as awhcle, tcccnstitutea surrcgate sludge with a prcpensityfcr phcsphcrus remcval, variaticninBEPR
betweendifferent systems was hypcthesited tc be due tc changes in the prcpensity fcr P remcv cf this
surrogate sIudge, caused by cnanges in inuent RBC0D ccncentration, anaercbic mass acticn and]cr
nitrate discharge tc the anaercbic reactcr. Bcwever, parIel research in the natur sciences had
identied specic crganis grcups that stcred pcIyphcsphate, this le1 tc a shift in the apprcach tc
cdelling BPR in NDBPR systems , frcm a surrcgate mass tc specic crganism grcups mediating
BPR, generically terme1 pclyP organisms [sc terme1 bioP crganisms [Ccmeau et al. , 1985);
reccmmen1e1 term, phcsphate accumulating crgamsms [PAO) by the IAWPRC Task 0rcup (1991)]. n
order tc 1evelcp the BPR kinetic behavicur cf NDBPR systems, attenticn ha1 tc be given tc the
icrobiclcgy an1biochemistry cfthesepclyP crganisms.
MICROBIOLOGY AND BIOCHEMISTRY OF POLYP ORGANISMS
Iuhs andChen[ 1975) ccnclude1 that the genus Acinetobacter was theprincipcrgamsmgrcup me1iating
BPR and thai "anaercbic ccnditicns preceding aercbicsis in sewage treatment cculd well be related tc
the appearance cf Acinetobacter". The presence cf Acinetobacter spp. in signicant ccncentraticns in
XDBPR systems was pcsitiveIy estabIished by Buchan [1981), Bart an1Melme1 (1982) , Buchan (1983)
and cthers.- Other genera have been impIicated in BPR, e.g. Pseudomonas (rc1isch and !cyner,
1983), but these have nctachievedsimiIar reccgnittcn,e.g. Wenttel et al. (1988a) in 1evelcpingenhance1
cultures cfpclyP crganisms [see later) fcund that when Pseudomonas spp. egantc 1cminate the culture,
Prelease, uptakeandremcvaldecIined signicantly.
Ircm the literature the fcllcwing characteristics cf the genus Acinetobacter were identied as cf
impcrtanceinBPR.
The genus requires cxygen as eIectrcn acceptcr [Juni 1978); hcwever scme species can use nitrate
when cxygen is nct present [Lctter, 1985). Mcst strns can grcw in a simple mineral medum
ccntaining a singIe carbcn an1 energy scurce such as ethancl, acetate, lactate, etc. Bergey, 1984);
relativelyfew strains can usegluccse as a carbcn scurcefcrgrcwth [Jum,1978), an1tenexclusively
via the ntner-Dcudcrcff pathway - Acinetobacter spp. 1c nct pcssess the mbden-Meyerhcf
[glycclytic) pathway [Juni , 1978). Acinetobacter spp. pcssess l cf the entymes cf the tricarbcylic
aci1 [TCA) cycle, as weIl as the entymes cf the glycxylate cycle [!uni , 1978). 5trns stcre
-Recently, the reliability cf the Analytic Prole In1ex [API) 2 i1enticaticn prccedure, ccmmcnly
use1 tc enumerate crganisms in activate1 sludge systems, has ccme un1er questicn - the prccedure has
been fcun1 tc cverestimate the Acinetobacter spp. ccunt [Venter et al. , 1989). Clearly, this aspect
requiresfurtherinvestigaticn.
6 M. C. WL 8lal.
(a) Htage Bardenpho system
[b) 5tage Mcdied Bardenpho system
(c) PC1 system
[e) 1chaaaeshutgsystem
[d) MPC1 system
hLX
ANAEROBIC
ANOXIC
OAEROBIC
Fg 1: 3chemat|c|aycutcvat|cus 3cuthAt|caa act|vateds|udgesystemccagutat|cas.
Fg 2:
50
40
:
..
,
30
0
C
,
a
20
;
0
1 0
0
(a)
O
0
I 1AWPHC UCTLLO X6flM6hI!
5 10 15
Time (h)
20 25
OPR-t|me prc|e |as|ag|ereactcr ccmp|ete|y aerc6|csystem at2.d s|udge ageuader da||ycyc||c
suare wave |cad|ag [eed I2h ca, I2h c, acte ptedp|tcus drcp |aOPR ca eed term|aat|ca.
OPR-t|me respcase s|mmated us|ag IAWPRC Task Grcup mcde| [IAWPRC), Heate et aI.
J)aadPCTmcde| [PCTOIB), Bc|d et a1. [1J).
NBEPR systs 65
carhca as pc|yhydrcxyhutyrate [PHB) aad phcsphcrus as pc|yphcsphate [pc|yP) [Fuhs aad Chea,
1975; Ue|aema et al. , 1980; Buchaa,1981).
0hservat|cascaXUBBPRsystemsgavethe!c||cw|ag:
0aer aaaerch|cccad|t|cas, shcrt cha !atty ac|ds [3CFA aad |atrace||u|ar pc|Pdecreased, sc|uh|e
phcsphate, Hg-, K* aad |atrace||u|ar PHB |acreased, aa

pH d|d act chaage Reas|ak, 1981; Hart


aa Hdmed, 1982; Fukase et al. , 1982; Wataaahe et aL, 1984; Arva, 1985; cmeau et aL, 1985;
Hasccet et aI., 1985; Weatte| et aI . , 1985; Hurphy aad Itter, 1986; Gerher et aL, 1987; Weatte|
et al. , 1988a}.
0aderaerch|cccad|t|cas, |atrace||u|ar pc|yP |acreased,sc|uh|ephcsphate,Hg-, K*aad|atrace||u|ar
PHB decreased, aad pH |acreased [Fukase et aL, 1982; Hart aad Hdmed, 1982; Wataaahe et aL,
1984; Arv|a, 1985; Ccmeau et aI . , 1985; Hasceet et aL, 1985; Hurphy aad Itter, 1986; Gerber et aL,
1987; Weatte| et aI . , 1988a) .
1e exp|a these chservat|cas a aumher c! mechaa|st|ch|cchem|c mcde|s were prcpcsed. The |mt|
mce|swere pr|adpa||y mechaa|st|c [Ha|| et aI. , 1978;

|chc||s aadOshcra, 1979; Reas|ak, 1981) aadthe


ma|aprcpcss caahesummar|tedas!e||cws:
(1) |crtcha|amttyac|s[3CFA)serveassuhstrate!crpc|yPcrgaa|sms.
[2) a the aaaerch|c phase the 3CFA are takea up aad stcred as PHB, th|s prccess requ|r|ag eaergy.
1he eaergy requ|remeats are supp||ed hy hreakdcwa c! pc|yP, w|th P he|ag rdeased tc the hu|k
se|ut|ca.
(3) Ia the aerch|c reactcr the stcred PHB |s used as a carhca aad eaergy scurce !cr cd| !uact|ca aad
gtcwth, aaasaa eaery scurce!crpc|yP !crmat|ca wh|chg|vesr|setcPuptake.
Marms et al. (1983) |atrcducedah|cchem|ca|Jmechaa|st|cmcde|hy putt|ag!crwarh|cchem|cpathways
er the syathes|sc

PHB uader aaaerch|cccad|t|cas, hewever, w|thacetate assuhstratethey!cuad|tact


pess|h|e tc |eat|Iy pathways !cr PHB syathes|s - ac scurce c! prctcas aad e|ectrcas, tc reduo
acetcacetate tc PHB, was ava||ah|e. 1heywere uaah|e, there!cre, tcadaa e|aaat|ca !crPHBstcrage
aaeraaaerch|cccad|t|cas w|thacetateassuhstrate. Ia 1982 F|creattaadHartemaaahadactedact|v|ty
of the tr|carhcxy||c ac|d [1CA cyde eatymes uader dch|c ccad|t|cas. Iadepeadeat|y, |a 1985,
Matsae [see H|ac et al. , 1987}
3
aad Ccmeau et aL prcpcsed that the prctcas aad e|ectrcas he
sapp||ed cm eperat|ca cI the TCA cyc|e uader aamrch|c ccadt|cas. Ccmeau et aL |deat|ed the
|mpcttaace c! mata|a|ag the prctca mct|ve !crce pm|) uader aaaerch|c cca|t|cas aad devdcped a
h|echem|c]ccaceptu mcd!cr BBPR. Hcwever, t

e|r mcde|descr|hedthe h|cchem|c pathwayscmy


|acut||ae. Weatte| et al. (1986) acceptedtheprcpcsa|s c! Ccmeau et aL thatuader aaaerch|c ccad|t|cas
t|e1CAcyde |s|avckedaad the pm! musthemata|aed,aadaccepted Acinetobacter spp.asthetyp|c
pe|yPergaa|sm med|at|ag8BPR. They prcpcseddeta||ed h|cchem|ca|pathways!cr Acinetobacter spp. |a
aaaerch|c aadaerch|creactcrsc!XUBBPRsystems,aadpreseateddescr|pt|cascIhcwthesepathwaysare
tegmatehy theA1P]AUP aad XAUH]XAU rat|es. Itter (1988) aad Itter aad Buhery (1987, 1989)
as|ag eatyme stud|es ver|ed the rega|atcry mechaa|sms aad that the TCA cyc|e ccu|d cperate uader
aaaetch|c ccad|t|eas. Hav|ag |deat|ed the regu|atcry mechaa|sms, Weatte| et aL (1986) app||ed the
h|cchem|c mcde| tc s|tuat|cas cther thaa the aaaerch|c]aerch|c XBBBPR systems, tc exp|a|a the
ehserved hehav|cur c! Acinetobacter spp.,suchass|mu|taaecusPHBaad pc|yP !crmat|ca |apurecu|tures
w|thh|ghacetateccaceatrat|casuaderaerch|c ccad|t|cas.
1he Weattd et al. mcde| was spec|c !cr Acinetobacter spp., |.e. !cr crgaa|sms not pcssess|ag the
Bmhea-HeyerhcI [BH) pathway, H|ac et aL (1987) prcpcsed a h|cchem|c mcdd!cr pc|yPcrgmsms
t|at pcssess the BH pathway, hut d|d act |

eat|

y the spedc pc|yP crgaa|sms. Th|s mcde| |s |a


agreemeat w|th that c! Weatte| et aL except that uader aaaerch|c ccad|t|cas the TCA cyc|e |s act
|mp||cated- prctcasaaddectrcasaresupp||edhyut|||tat|cac! stcredcarhchydratev|atheEHpathway.
M|ae et aL vet|ed exper|meata||y the ut|||tat|ca c! stcred carhchydrate uader aaaerch|c ccad|t|cas,
at|||tat|ca c! eztraceUular carhchydrate uader aamrch|c ccad|t|cas has heea chserved per|meat|y hy
0shera et aL (1989). Weatte| et al. (1991) adaptedthe H|ac et al. mcde|!cr crgaa|sms thatpcssessthe
Bataer-Ucudcrc [

U)pathway, |a th|sadaptedmcde| the TCA cyde sc |sact|mp||cated- prctcas


aaddectrcasare supp||edhyut|||tat|ca c!stcredcarhchydratev|athe EBpathway.
ircm aa evuat|ca c! the prcpcsed h|cchem|c mcde|s, CcmeauJWeatte|

H|ac aad adapted H|ac,


weattd et al. (1991) ccac|uded that there |s ccaseasus regard|

maay 01 the h|cchem|c pathways


|avc|ved, hut that the scurce c! reduc|ag equ|va|eats ABH reqmred |a the reduct|ca step |a
ccavert|a

acetate tc PHB rema|as tc he estah||shed, |.e. whether the NABH |s supp||ed v|a the TCA
[CcmeauWeattd), BH [H|ac) cr BB [adapted H|ac) pathways, they prcpcsed aa apprcach tc resc|ve
Hatsuc's(1985) paper|s|a1apaaeseaadccmdacthestumed!crtmsrev|ew.
6 M. C. WZL 0lal.
this questicn but this stu1y hasnct yet been un1ertaken. 0neimpcrtant ccnc|usicn was that irrespective
cf whichbicchcmica| mc1e is cperative, thepclyP crganisms will be unable tcutilitegluccse1irectlyfcr
PBB prc1ucticn un1er anaercbic ccn1iticns. Acccr1ingly they ccnclu1e1 that the prcpcss cfWenttel
et al. [ I) remain va|i1 - that the gluccse-like materia| present in the RB00D acticn cf 1cmestic
wastewater requires ccnversicn by ncn-pclyP crganisms tc 50IA un1er anaercbic ccn1iticns befcre
utilitaticn by pclyP crganisms. The impcrtance cf this ccnclusicn w|ll beccme apparent in mc1elling
BPRinmixe1 cmture systems, seelater.
Baving largely resclve1 the bicchemistry]micrcbiclcgy, attenticn ccul1 be fccuse1 cn the kinetics cf
NDBPRsystems.
NDBEPR SYSTEM KINETICS
Wentte et al. [ Ia) set cut tc 1evelcp a gener kinet|c mc1el that 1escribes XDBPR system
behavicur. They assume1 that in a NDBPR system treating municip wastewaters a mixe1 cmture
wcm1 1evelcp which ccul1 becategcrite1intc threegrcups cfcrganisms [ I) heterctrcphiccrganisms able
tc accumu|ate pclyP, terme1 pclyP crganisms, [2) heterctrcphic crganisms unable tc accumulate pclyP,
terme1 ncn-pclyP crganisms, an1 [l) autctrcphiccrganisms me1iating nitricaticn, terme1 autctrcphs.
Wenttel et al. reccgnite1that 1evelcpment cf an activate1slu1gek|neticmc1eltc1escribethe behavicur
cf NDBPR systems wcul1 require inclusicn cf all three crgamsm grcups, an1 their interacticns. With
regar1 tc ncn-pclyP an1 autctrcph|c crganisms, they accepte1 the general ND kinetic mc1el 1escribe1
earlier[Dcl1 et al., I, van Baan1el et aL, II, Bente et al., II, Dcl1 et aL, II). Thismc1elncw
nee1e1 tc be exten1e1 tc inccrpcrate pclyP crganism behav|cur in cr1er tc 1evelcp a mc1el that wcU1
inc|u1e l three crganism grcups - a general NDBPR kinet|c mc1el. Tc achieve this cbjective, the
kinetic an1 stcichicmetric characteristics cf the pclyP crganisms in te avaed sludge envrnmen
nee1e1tcbeestablishe1.
Ircm attempts tc cbtain infcrmaticn cn the characteristics cf the pclyP crganisms using mixe1 |iqucr
cm XDBPR systems treating municipal wastewaters, Wenttel et aL [1a) ccnclu1e1 that, in these
m|xe1 culture systems, the ncn-pc|yP crganism behavicur ten1s tc 1cminate an1 mask the pclyP
crganism behavicur. Acccr1ingly, they en1eavcure1 tc isclate the pclyP crganism characteristics, by
1evecping enhanced cultures c these crganisms in activate1 slu1ge systems. By enhance1 cu|ture is
meant: A cu|turein which the grcwthcf pclyP crganismsisfavcure1 tc the extent that they beccme the
1cminant primary crganism an1 their behavicur 1cminates the system respcnse, grcwth cf ccmpeting
crganismsiscurta|le1 but nct pcsitively eclu1e1, neither arepre1aticncr cther interactive effects, sc,
astrn]spec|e[s)cf pclyP crganismisnaturly selecte1intcthesystem. Wenttel et al. [ Ia) prcpcse1
tcachieveapclyPcrganismenhance1culturebyselecting asubstratean1 set cf envircnmentalccn1iticns
inanactivate1slu1ge system thatwcul1greatlyfavcurpclyP crganismgrcwth.
ENHANCED POLYP ORGANSM CULTURES
Enc ctue deveopment
Ircm the bicchemica| mc1els, Wenttel et al. [Ia) were able tci1entify ccn1iticns tcbe impcse1 in a
NDBPR activate1 slu1ge system tc prc1uce an enhance1 culture - anaercbic]aercbic sequence with
a1equateanaercbicmassracticn inuent fe1 tctheanaercbicreactcrwithacetateas substratean1with
a1equatemacrc- an1 micrcnutrients, inparticular Hg-', K'an1 tc a lesser 1egree 0a-' ; ad pB ccntrcl
intheaercbicreactcr. 0sing the 00T an1 l-stage Hc1ie1 Bar1enphc systems, withsystem slu1geages
ranging frcm I. - 21, they 1evelcpe1 enhance1 cultures cf the pclyP crganism Acinetobacter spp. -
>% cf the crganisms culture1 aercbically were i1entie1 tc be Acinetobacter spp. using the Analytic
Prcle In1ex [API) 2 prcce1ure. The respcnse cf the enhance1 culture systems in1icate1 that
s|gnicant ccncentraticns cf pclyP crganisms 1evecpe1, fcr example

the 00T system [anaercbic mass


fracticn I%, slu1ge age I1an1 inuent cf acetate at mg00D] gave P release 2l, P uptake
3I1 an1 P remcva| I, all mgP]l inuent cw [cf. mixe1 culture NDBPR systems with munidp
wastewaterinuent cf mg00D]( P release 1, P uptake I, an1 P remcv I2 mgP]linuent
cw). Theenhance1 culture mixe1liqucrintheaercbictcneccntaine1 . lmgP]mgHLV55an1ha1a
V55]T55 ratic . 1 mgV55]mgT55 [cf. P]HLV55 . I an1 V55]T55 . I fcr mixe1 culture
systems). The lcw V55]T55 raticfcr theenhance1 culture systems was 1ue tc the massive amcunts cf
pclyP withasscciate1ccuntericns [principly Hg-' .2mcl Hg-']mcl P an1 K' . lmcl K']mcl P)
stcre1bythepclyP crganisms.
NBEPR systems
67
Enhace cutu knetic mode
Pto experiment obsetvations on the enhance1 cultute stea1y state systes, an1 on batch tests in
wh|chixe1 liquots 1tawnftothe stea1y state systes wetesubjecte1to a wi1e vatiety ofcon1itions,
Wentte et al. [ Ia) euci1ate1 the charactetistics an1 kinetic tesponse of the polyPotganism mass.
Twochatactetistics cfthepolyPotganismsintheenhance1cultutesweteofparticularintetest.
[I) Littleptopensity to 1enittify so that no ptovision fot this ptocess n1e1 to b ma1e in mo1elling
polyP otganis behaviout. [ The lack of 1enittication by the polyP otganiss has iplications in
o1ellingixe1cultuteNDBPRsystes, seelatet) .
[2) An extteely low en1ogenous ass loss tate, . 1 gAV55][mgAV55. 1) cf. the en1ogenous mass
loss tate of . 21 mgAV55][mgAV55. 1) 1etet|ne1 fot aetobic activate1 slu1ge systes [ie.
non-polyP otganiss) by HcKinney an1 0oten [ I) an1 Hatais an1 kaa [II)}. A similat
obsetvation ha1 been a1e by Wenttel et aL [ I) in stu1ies on mixed culture NDBPR systes
tteatingunicipalwastewatets, they note1 fto plots ofP uptakevetsus P teleasefotvatiousslu1ge
agesthat,fot agivenP telease_theP uptakewastelativelyinsensitiveto slu1geage. Toexplainthis
obsetvation, Wentte| et al. I) ha1 ptopose1 that "the polyP otganiss suet little ot no
en1ogenous ass loss". The high specic en1ogenous mass loss tate with non-polyP otganism
systes ha1 been atttibute1 to a high tate of pte1ation an1 tegtowth, fotulate1 as 1eath
tegenetation inthe ND kinetic mo1el by Dol1 et al. , I [see eatliet) . The low specic en1ogenous
ass loss tate with polyP otganiss in enhance1 cultute systes an1 the obsetvations of Wenttel
et al. [ I)le1Wenttel et al. [ Ia)toconclu1ethatpolyP otganisms ate not pte1ate1 to the sae
1egtee as non-polyP otganiss. Accot1|ngly, in o1elling polyP otganism en1ogenous mass loss,
Wentte et al. use1theclassical en1ogenous tespitation ap

toach, except thatptovsionwasma1efot


situations wheteno extetnalelecttonacceptot is availableseelatet).
Taking note of the above, Wenttel et al. [ Ia 1evelope1 a conceptual o1el fot polyP otgams
behaviout intheenhance1cultutes incotpotating te chatactetistics , ptocesses an1 copoun1s i1entie1
as ipottantfto the expetient investigation. 0singtheconceptu o1el as a basis, Wenttel et al.
[Ib) fotulate1 mathematicly the ptocess tates an1 theit stoichioetric intetactions with the
copoun1s, to 1evelop a kinetic mo1el fot the enhanced cultures of polyP organisms. This mo1el is
ptesente1 inTable I using the attix fotat tecoen1e1 by the IA WPRC Task 0toup [Bente et aL,
II). 0sing the enhance1 cultutes, thekinetic an1 stoichioettic constants wete quantie1 byavariety
ofexperiental ptoce1utes [Wenttel et al. , Ib).
With these constants, application ofthe kinetic mo1el to the various test tesponses obsetve1 with the
enhance1 cultutes gave goo1 cottelation between obsetvations an1 siulations [see Pigs l to ) . The
o1el then was applie1to simulatethe stea1y statebehaviout oftheenhance1cultute 0CT an1 ltage
Mode1Bat1enpho systes, againgoo1cottelationwasobtne1,Wenttel et al. [ Ib) .
Simplfed enhanced cutue steady state mode
Wenttel et al. [I) simplie1 the enhance1 cultute kinetic o1el, to 1evelop a stea1y state mo1el fot
the enhance1 cultute systems un1et constant ow an1 loa1 con1itions - ftom an examination of the
kinetics ofthe ptocesses un1et stea1y state con1itions, they foun1 many ofthe ptocesses to be vittually
coplete, these kinefic telationships no longet setve1 any function an1 coul1 be teplace1 by
stoichioettic telationships. Putthet, they ma1e two assumptions which simplie1 1evelopment ofthe
stea1ystatemo1e.
[I) P telease fot anaetobic maintenance enetgytequtements is ways smallcompate1 to P te|easefot
sequesttationenetgytequteents, ie. the Hnetics of Ptdeasefot anaetobicmaintenanceenetgynee1
not be incotpotate1. The implication of this is that un1et stea1y state the polyP content of the
polyPotganissinthe activate1 slu1ge waste1 pet 1ay,isconstant.
[2) Al the substtate sequestete1 in the anaetobic toneisutilite1 inthesubsequent aetobic tone, ie. the
kineticsofPBB substtateutilitatlonnee1notbeincotpotate1.
Taking 1ueaccount ofthese assuptions , an1 theobsetvationthat theanaetobic assactionsptovi1e1
intheenhance1 cultute systes were sumcientto ensutethat all theacetate substtate was sequestete1 in
theanaetobictone [ie. kineticsofacetatesequesttation nee1 not be inclu1e1) , Wenttel et al. 1evelope1 a
nubet of stea1y state equations fot the enhance1 cnltutes, fot , polyP otganism active an1 en1ogenous
asses, an1 P telease, uptake an1teoval 1ue to these masses. These equations ptovi1e1 the means fot
uantifying the polyP otganism population in ixe1 cultute NDBPR systes teceiving municipal
wastewatets asinuent, seebelow.
T
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N8EPRsysts 0
.1 m1-
1"1
4 5 ro
TIME (day.)
Fg 3(a): Experimentaly observed and simulated
oxygen utilization rate (OUR) response with time
in a batch aerobic digestion of mixed liquor fom
the enhanced culture system (Wentzel 6l 0
1989b).
DD
TIME (days)
Fg 3(e): Experimentaly observed and simulated
fltered COD concentration-time profles for the
batch aerobic digestion in Fig 3(a).

TIME (I.)
F . Experimentaly observed and simulated
tota soluble phosphate concentrations (POc) and
carbonaceous oxygen utilization rate (Oul) on
aeration following aaerobic acetate addition of
0. mgCOD acetate/mgVSS to a mixed liquor
batch fom the enhaced cuture system (Wentze
6l 0 1989b).

i
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.

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a
..
2
B
.
ID e
U
D
O
TIME (day.)
PO
- 11 m1a
E1-1
r
Fg 3(b): Experimentalr observed and simulated
tota soluble phosphate (POc) and nitrate (N03)
concentration-time profes for the batch aerobic
digestion in Fig 3(a).
IOrrIO
.
-Ss_lal

r .
Exl.l
I.

TIME (hrs)
F 4: Experimentaly observed ad simulated
total soluble phosphate (POe) and acetate
concentration-time profles Wth anaerobic addition
of 0.11 mgCOD acetate/mgVSS to a mixed liquor
batch fom the enhanced culture system (Wetze
6l 0 1989b).
a
u
5


D
O I.
TIME (hr.)
- ., . Ia
p ..l
II ta
F . Experimentaly observed and simulated
total soluble phosphate concentrations (POc) ad
carbonaceous oxygen utilization rate (Oul) on
aration following aaerobic acetate addition of
0.ZZ mgCOD acetate/mgVSS to a mixed liquor
batch fom the enhanced culture system. The POc
concentration falls to zero during the course 0 ths
test (Wentze 6l 0 1989b).
70 M. C. WL 8lal.
STEADY STATE MIED CULTUR NDBEPR SYSTEMS
Me cutu stedy statemMd
Bav|ag deve|cped the steady state mcde| !cr eahaaced cu|ture systems, Weattd et al. (1c) exteaded
th|smcde| tc|accrpcratem|xed cu|tures c! pc|yPaad aca-pc|yP crgaa|sms preseat|aN

BEPRsystems
tece|v|ag dcmest|c wastewater as |a0ueat, tc g|ve a steady state m|xed cu|ture mcde|. Th|s exteas|ca
prcvedpcss|h|ehecauseeahaacedcu|turesratherthaapurecu|tureswereused tcestah||shthek|aet|caad
stc|ch|cmetr|ccharacter|st|csc!thepc|yP crgaa|sms. Ia the eahaaced cu|tures, stras [spec|es) c!pc|yP
ctgaa|sms preseat |a m|xed cu|ture act|vated s|udge were ear|ched [us|ag uaderstaad|ag c! the
h|cchem|stryaadm|crch|c|cgyc!pc|yPcrgaa|sms)aadas|ag|estra [spec|e)wasactart|c||y se|ected
[as |a

ure cmtures), ccmpet|ag crgamsms aad predatcrs were act art|

c|a||y c|uded [as |a pure


cu|tutes) sc that the pc|yP crgaa|sms were suhjected tc the same se|ect|ve pressures |a eahaaced as |a
mxedcu|tures, thepc|yPcrgaa|sms a|sc weresuhjected tc the same ccad|t|cas

reseat |am|xedcu|tute
act|vated s|udge systems [e.g. aaaerch|c]aerch|c sequeac|ag, |cag R > d, etc.), the pc|yP crgaa|sms
h|h|ted the same hehav|cur patteras |a the eahaaced cu|tures as they dc |a m|xed cu|ture act|vated
s|udge systems [ie. P tdease]uptake, PHB]pc|yP accumu|at|ca, etc.) - |a !act, the s|m||ar, thcugh
'maga|ed' hehav|cut c! the eahaaced cmture ccmpared tc the m|xed cu|ture systems was cae cr|ter|ca
used tc estah||sh that the ccrrect eahaaced cu|tures had hea estah||shed. Ia extead|ag the mcde| cae
upect that emerged was the d|!!ereace |a the eadcgeacus mass |css rate hetweea pa|yP crgaa|sm
eahaaced cu|ture s|udges aad the "acrm" aerch|c aca-pc|yP crgaa|sm act|vated s|udge [see sect|ca
'@ cutu knc mMd"). As acted ear||er, the h|gh spedc eadcgeacus mass |css rate w|th
aca-pc|yPcrgaa|sm systems hadheea attr|hutedtc a h|ghratec! predat|caaadtegrcwth, !crmu|ated as
death regeaerat|ca |athe NBk|aet|cmcde| hyBc|d et al. [Ic). 1he|cwspedceadcgeacusmass|css
tate w|th pc|yP ctgaa|sms |a the eahaaced cu|tures systems had |ed Weattd et al. [Ia) tc ccac|ude
thatthepc|yP ctgaa|sms were actpredatedtc thesame degreeasaca-pc|yPcrgaa|sms, aadtcadcptaa
eadcgeacusresp|rat|caapprcach|amcde|||agpc|yPcrgaa|smeadcgeacus mass |css. The|cw predat|ca
tate ca the pc|yP crgaa|sms, aad the !act that the pc|yP aad aca-pc|yP crgaa|sms esseat||y dc act
ccmpete cr the same suhstrate, |mp||ed that pc|yP aad aca-pc|yP crgaa|sm pcpu|at|cas act v|rtua||y
|adepeadeat|yc!eachcther|a"acrm"m|xedcu|tureNBBEPRsystems. Iadeve|cp|agthesteadystate
mcde|ct m|xed cu|ture NBBBPR systems, Weatte| et aL [Ic) acted that th|s |mp||es that aays|s c!
the twcpcpu|at|cagtcupscaahe|argdyseparated. Hcwever, twcs|ga|caat |ateract|caswere|deat|ed
ct|ac|us|ca|athem|xedcu|tureNBBEPR steadystatemcde|, hcth|athe aaaerch|c reactcr, as!c||cws:
[I) I maay "actma|' mua|c|pa| wastewaters the acetate [cr cther shctt cha atty add [3CFA)
ccateat|s sma|| ct act preseat [Weatte| et al., Ih). Weatte| et aL [I) had shcwathat |a t

e
aaaetch|c teactct the RBCOB ccmpcaeat c! the |a

ueat |s ccaverted tc 3CFA hy the aca-pc|yP


ctgaa|sms, thetehy mak|ag 3CFA av|ah|e tcthe pc|yPcrgamsmmass!crsequestrat|ca. Thetatec!
ccavets|ca |s much s|cwer thaa the rate c sequestrat|ca, sc that the rate c ccavets|ca ccattc|s the
tate c sequestrat|ca. Beace, the mass c! 3CFA suhstrate that heccmes ava||ah|e |a the aaaetch|c
teactct tcthe pc|yP crgaa|sms |s gcveraed hy the k|aet|cs c! ccavers|ca med|ated hy the aca-pc|yP
ctgaa|sms. The wcrk c! Hegaack et al. [I) aad Brcd|sch [I) suppctts th|s ccavets|ca
hypcthes|s, they had shcwa that aaaerch|c]aerch|c systems deve|cp crgaa|sms wmch ccavert sugars,
ds|m||mccmpcuads, tc3CFA|atheaaaerch|creactcr.|
(2) I mtrate [cr cxygea) |s recyc|ed tc the aaaerch|c reactcr, RBCOB |s ut|||ted prdereat|m y hy the
aca-pc|yPctgaa|sms w|tha|trate[crcgea)as exterae|ectrcaacceptcrtherehyreduc|agthemass
cRBCOD ccavertedtc3CFA.
Weattd et a [I)had recga|ted [I) aad (2) ahcve, aad !crmu|ated ak|aet|c mcde| !crccavers|cac!
RCOD tc 3CFA, aad heace !cr sequestrat|ca c! these 3CFAs. Weatte| et al. [Ic) accepted th|s
ccavets|ca mcde|, hut made prcv|s|ca tc |ac|ude s|tuat|cas where 3CFA are preseat |a the |a0ueat hy
act|ag that thetate c! 3CFA sequestrat|ca |s scrap|dthat all infuent SCFA w||| he sequestered hy the
pc|yP ctgaa|sms |a theaaaerch|creactcr!cr aaaerch|c mass act|cas >IcX aad s|udgeages >I0d [th|s
caa b vet|ed rcm the k|aet|cs c! sequestrat|ca). Th|s thmry prcv|ded Weatte| et aL [Ic) w|th the
meaas ct ca|cu|at|ag the mass c! 3CFA suhstrate [cm the |a0ueat aad cm ccavers|ca c

RBCOB)
sequestered hy thepc|yPcrgaa|sms|a theaaaerch|creactcr. Kacw|ag the mass c! suhstrate sequestered
6y thepc|yP ctgaa|sms, themassc!suhstraterema|ag, ava||ah|etctheaca-pc|yP crgaa|sms, ccu|d he
cmcu|ated. Ia e6ect Weatte| et aL sp||t the h|edegradah|e |a0ueat COB |atc twc !ract|cas, cae
eveatum|y tc6eut|||tedhythe pc|yP ctgaa|smsaad the cther tche ut|||ted hytheaca-pc|yPcrgaa|sms.
8eausecthe|adepeadeacec!act|cac!thesetwcgrcupsc!crgaa|sms, theyccu|duse:
Frcms|mu|at|cassu6sequeat|yw|ththesteadystatem|xedcu|turemcdd,|twascuadthat, |!thepc|yP
ctg|sms wetesuhjectedtca h|ghptedat|carate, theas|ga|caatBEPR|a them|xed cu|tute NBBEPR
system wcu|d act he pcss|h|e - the tate c! death c! the pc|yP ctgaa|sms wcu|d he sc h|gh that ac
signicat mof tese organisms ccu|daccumu|ate|athesystem,aadBEPRwcu|dheaearterc.
NBEPR systms
71
[I) The simplie1 polyP otganism enhance1 cultute stea1y state mo1el fot calculating the polyP
organism active an1 en1ogenous masses fotme1 ftom the sequestete1 substtate, an1 the P telease,
uptakean1temoval me1iate1 bythesemasses.
(2) The stea1y state activate1 slu1ge mo1el [Mats an1 kama, II, WRC, I1) to cculate the
aoa-polyP otganism active an1 en1ogenous masses fotme1 om the temning substtate, the rate of
coaversioa of RBC0D to 5CPA in the anaetobic reactot, the inett V55 accumUate1 from the
imueat, an1 the P requitement of, an1 hence P temov associate1 with, the active, en1ogenous an1
iaert masses. (Notethatiathisstea1ystateactivate1slu1gemo1elen1ogenous mass lossismo1ele1
using theclassical en1ogenous tespitation apptoach - this apptoach is simplet an1un1er stea1y state
coa1itionsgivesresUtsvetydosetothe1eathregenetationapptoach) .
The tot P removfotthesystemwascculate1 bysummationofthein1ivi1uP removs.
Wentte et al. [I) evuate1 the pte1ictive powet of the stea1y state mixe1 cultute BPR mo1el
agast observations ma1e on l labotatoty sce NDBPR systems over a six year perio1, system
coagurations were Phote1ox, ltage Mo1ie1 Bat1enpho, 0CT, H0CT an1!ohaaaesburgwith system
slu1ge ages rangiag om l to 2 1ays. Pot the evuation the measute1 nittate ia the recycle to the
aetobic tone was use1 to estimate the RBC0D temov in the anaetobic tone by the non-polyP
organisms with aitrateas extern electton acceptot. TheRBC0Dremningwas avlablefor convetsion
iatheanaetobicreactotto 5CPA, fotsequesttationby thepolyP organisms. Plotsofthe pre1icte1 vetsus
measure1P release, P temoval an1 V55 concenttation, Pigs I to , show goo1cottdation.
Icrpration o dentrfcation apcts in steady state me cuture mode
Ia the stea1y state phosphorus evuations using the mixe1 cultute mo1el [ Pigs I to ) , necessarily the
aitrate recyde1 to the anaetobic teactot nee1e1 to be known, an1 this was avlable om expetiment
obsetvations on the NDBPR systems. Cleatly fot completeness 1enittication ha1 to be incorpotate1
iato the stea1y state mixe1 cultute mo1el, an aspect omitte1 up to this stage. 0ne possibility to
accomplish thiswas toestimatethenittateinthetecycleto theanaetobicteactotomthe1enitricatioa
theoryfor the ND stea1y state mo1el [kama et al. , Il, WRC, I1) . xpetiment 1ata in1icate1
thatthe ND stea1y state mo1eptedcte1the1enitticationinNDBPRsystems quitedosely. Bowever,
withthe1evdopment ofthe BPRtheoty, in applying the ND stea1y state mo1e to NDBPRsystems
aaiacoasistencyintheapptoachbecameevi1ent.
The eahaace1 culture stu1ies in1icate1that polyP otganisms 1o not 1enittify. This implie1 that the
R00D, coaverte1to 5CPAbythe non-polyP otganisms an1 sequestete1bythepolyP orgisms ia
the aaaerobic reactor, ao longet was avlable fot 1enittication in the ptimatyanoxcreactorof a
NDBPR system. This ia turn implie1 that the magnitu1e of the 1enitrication ia the primary
aaoxcreactoroftheNDBPRsystem shoU1 be signicantly smlerthaathatiathepnmary aaoxc
reactor ofthe ND system. Bowever, expetimeat observations on NDBPRsystems ia1icate1 that
thiswasaot so, that approxmately the same magnitu1eof1enitticationwasachieve1. Theimplica-
tioa was that the 1enittication kinetics fot ND systems nee1e1 to be a1apte1, or mo1ie1, fot
applicatioaia NDBPRsystems.
0siag plugow aaoxic teactots an1 batch tests, Clayton et al. [ I, II) un1ettook an

xpetimeatal
iavestigatioa into the kinetics of 1enittication ia NDBPR systems. hey foun1 that :n NDBPR
systems,
[I) iatheprimatyaaoxic reactor, (a) the rapi1 rateof1eaitrication associate1 with RBC0Dwas much
re1uo1orabsent, (b) theslowet rateof1enittication associate1 with 5BC0Dwas approxmate 2
timestheratemeasure1inptimatyanoxicteactotsofND systems, an1
(2) iathesecoa1aryaaoxicreactor,the1enitticationtatewas apptoximately I times theratemeasute1
iasecon1aty anoxcreactotsofNDsystems.
Prom aa extensiveenquity into causes, Clayton et al. condu1e1 that the inctease1 1enittication tates
were not 1ueto,
[ I) deaitricationbypolyPotganisms- fot thesystemsinvestigate1, PBBan1P measutementsin1icate1
thatthepolyPorganisms1i1not 1enittify,
(2) mo1icatioaofthe sewage ia the anaetobic tone- sewage that ha1 not passe1 thtough an a
.
naetobic
toneia1uce1 thesame1enitticationtesponseassewagethat ha1passe1 thtough theanaetob:c tone.
The above observations le1 Clayton et al. to condu1e that the inctease1 rate is 1ue to a stimulatioa ia
theactiveslu1gemassofaninctease1tateofhy1tolysisof5
.
BC0Binhe anoxicteactorsof

heNDBP
.
R
systems appateatly ia1uce1 by the presence ofthe anaetob:c teactot :n thesesystems. (Tks aspect st:ll
nee1s t beincotpotate1quantitativelyinto thestea1ystatemixe1 cultute mo1el, but this isnotexpecte1
topreseatua1ue1imculties. )
72 M. C. W 8l al.
g 8. Pre1icte1 versns easnte1 P reoval,
pre1ictions nsing the ixe1 cnltnre stea1y state
o1el, 1ata o labotatory scale systes
[Wenttel et e| , I).

a Id

+ 4
O O

V b
V
0 1 O


O
b
400D
Fg 7: Pte1icte1 versns easnre1 P telease,
pte1ictions nsing the ixe1 cnltnre stea1y state
o1el, 1ata o laboratory scale systes
[Wenttelet e| , I).

e
~
2
20
g
f "

D
f lD
l
L
~

a Id1
d
4
O
O
V b
Y
1 O
U

O
b

g 9. Pre1icted versns easnre1 V55


concenttations, pre1ictions nsing theixe1 cnltnre
stea1y state o1e, 1ata o laboratory sce
systes [Wentte et e|. , I) .
NBEPR systs 73
GENERAL NDBEPR KNETC MODEL
The stea1y state mixe1 cnltnte BPR mc1el, 1esctibe1 abcve, was testticte1 tc ccnstant cw an1 lca1
con1itions. Tms testticticn ma1e it pcssible tc accept cettain behavicnt pattetns which latgely
eiinate1 the nee1 fct ccmplete kinetic 1esctipticns cf the ptccesses. [Pct example, acceptance that
PHB is ccpletely ntilite1 in the aetcbic teactct eiminate1 the n1fct a 1esctipticn cfthe kinetics cf
PBB utilitaticn, an1cthets. ) Tc 1evelcp a mc1el that 1esctibes NDBPRsystem behavicnt nn1etcyclic
ow an1 loa1 con1iticns, snch simplicaticns nc lcnget ccnl1 be accepte1 - this teqnite1 the integtaticn
otheenhance1cnltnte BPRkinetic mc1elwiththegenetactivate1sln1ge ND kinetic mc1el tcgive a
gener activate1 sln1ge NDBPR kinetic mc1el, taking 1ne acccnnt cf any intetacticns between the
popnlaticn gtcnps inttc1nce1 by the integtaticn. This mc1el is neating ccmpleticn, its ptesent state cf
1evecpment is givenbelcw.
hDBBPRHaeicmodddesmgtioa
Pct the integtate1 genet activate1 sln1ge NDBPR kinetic mc1e, the ptccesses, thmt kinetic rate
ctUaticns an1 theit stcichicmettic intetacticns with the ccmpcnn1s ate set cnt in mattix fctmat in
Table 2. Pteliinaty estimates cf vnes fct the stcichicmettic an1 kinetic ccnstants in the mc1e ate
liste1inTable l[a,ban1c). 5ymbcl system nse1 is liste1 inTable 1.
Referting tc Table 2, the genetal NDBPR kinetic mc1e incln1es 2 ptccesses an1 I ccmpcnn1s.
PtccessesI-I2 an1ptccess2teectncn-pclyPctganismbehavicnt,ptccesses Il-I1antctrcphicctganism
behavicnt an1ptccesses I-21pclyP ctganismbehavicnt.
With tegar1 tc the aoa-pclyP aa1 autotrcphic organisms, the prccesses [prccesses I-I1), thmr rate
ctulaticns an1 stcichiometric intetacticas with theccmpcunds remn the same as fcrthe ND Haetic
c1el, 0mvetsity cf 0ape Towa versica [Dol1 et al. , II, i. e. death regenetaticn-bisubstrate syathesis
mc1el,with theexcepticathat theinteracticnsbetweeatheseprccesses an1theccmpcun1phcsphorusare
a11e1. Ptcmimtial simnlaticns [seelater), it wcU1appeatthat the asscoate1 kinetic an1stoichiometric
ccnstants alsc remain the same as fcr the ND mo1el, with one exception - the value fcr ' whichis the
taticcf the hy1tclysis]ntilitatica ratefcr 5B00D un1er anoxc aad aetcbic ccnditicns, i . e. 'G = aaoxc
tate[aetcbic rate. The vue for 'G had to be iacreased, bcm 0. ll to approxmately 0. to rect the
epetimentally cbserve1 1enitticatica by acn-pclyP otganisms ia NDBBPR systems [s earlier) .
5imnlaticns nsing the NDBPRkiaetic mo1el with the iacrease1 vue fot 'G shcwed that theiacreased
1enitticaticn rates ia primary aad seccndary aaoxc reactors ofNDBPR systems are, as aoted earlier,
abcut2times an1 I times respective|y theratesforhDsystems.
In a11iticn tc ptccesses cbtne1 bcm the ND kinetic mc1e, an extraptccess me1iate1 by aca_olyP
ctganisms (i.e. process 25), conversion ofRB00D to 50P A un1er aaaercbic ccn1itioas is iacluded ia the
NDBPR kinetic mc1e, to expaa1 1escription of ncn-pclyP ctganism behavionr un1er anaerobic
ccn1iticns. Pctmnlaticn cfthis prccess rate an1 stcichicmetry was taken nnmc1ie1 bom the eahaaced
cnltute kinetic mc1el whete it was incln1e1 even thcngh it serve1 nc kinetic fnnctica) for the geaer
NDBPRkineticmc1.
With tegat1 tc the pclyP ctgamsm 1esctipticn [ptccesses I-21), the ptccesses, the process rate
ctmnlaticns an1 kinetic constants remain thesame as intheenhanced cnltnte kinetic mcdel. Aspects of
thissecticn cfthemc1el that wattantmenticn, ate.
[I) Inline with expetiment cbsetvaticns [Wenttel et aI. , Ib) , pclyP ctganisms seqnester 50PA aa1
stcte these as PBB ua1er anaetcbic, ancxc an1 aetcbic ccn1iticns [ptocess 21), hcwever, ia
applicaticn cf the mc1e tc NDBPR systems this ptccess will be cpetative cnly nn1er anaercbic
ccn1iticns becanse signicant 5CPA ccncenttaticns ate likely tc be enccnntere1 cmy nn1er these
ccn1iticns. Te rtes cf 50PA seqnesttaticn an1 PBB stctage atemc1elle1 as in1epen1ent cf 50PA
ccncenttaticn an1 tst ct1et with tespect tc pclyP ctganism biclcgic active mass ccncenttatica
[ftcm eperimental cbsetvaticn, Wenttel et al. , Ib). 0cnccmitant tc 50PA seqnesttaticn, stcre1
pclyP iscleave1 and P telease1 tc thebnlk sclnticn. ThepclyP cleavage an1 P telease ate mo1elle1
as being 1irectlyptcpctticn tcthe50PAseqnesticn, d cbsetve1expetimently.
[2) PclyP ctganisms gtcw cnly nn1et aetcbic ccn1iticns, an1 cmy with stcte1 PBB d snbsttate
[ptccesses I-I). As PBB is an intetnl stcte1 pclymet [i. e. patticnlate) , the specic rate cf
ntilitaticn [an1gtcwth) is mc1ele1 as a sntface satntaticntypeteacticn similat tc that propose1fcr
5B00D hy1tclysis]ntilitaticn [Dcl1 et aI. , I0, see eatliet) . Ditectly ptcpctticn tc PBB utilited,
P is taken np fct pclyP fctmaticn an1fctcel synthesis [as cbsetve1 expetimently, Weattel et aI. ,
Ib), shcnl1 sclnbleP beccmelimitiag, P nptakefcrpclyP fctmaticnceases, bnt grcwth coatiaues
with the P tequitements for cdl syathesis snpplie1 om the pclyP pccl. Por growth, ammoaia is
use1 as a nittcen scutce, shcnld ammonia become limiting, nittate can serve as aa teraative
nittcgen scntce ct gtcwth. Pcllcwing the expetiment cbsetvaticns cf Wentte et al. [ Ib) aad
74
Table 2:
M. C. 8lal
Prccess Hnetics an1stcichicmetryfcrnitricaticn 1enitricatica biclcgicu ecesP temcvuin
NDBPRsystems.
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M. C. WENTEL 0l al.
0|aytca et al. [I) pc|yP crgaaiss canact use nitrate as exteraa| electrca acceptcr, i . e. cannct
grcwun1er ancxic ccn1iticns aa1acccr1iag|y "perceive"ancxic cca1iticasas anaercbic.
[l) A c1ie1 ea1cgencus respiraticn apprcach is use1 tc 1eal with

clyP crgani sm en1ogenous mass


lcss. n1c

eacus mass |css i s c1e||e1 separate|y fcr theaercbic prccess I) an1 anaerobic]ancxic
[prccess 2 states 0a1er aercbic ccn1iticns , a fraction of the biolcgic active mass lcss
[bic1egra1able pcrtica is |ccate1 tc an cxyen 1ean1, tc acccunt fcr maintenance energy
requireeats , aa1 the a|aace a||ccate1 tc sce cr cfiacrt ateria|, thetcta| being equa| tc the
"active ass |css rate", the active ass loss rate is c1e|le1 as prcpcrticnal to the active mass
ccnceatraticn 0n1er anaercbic an1 anoxic con1iticns , because no suitableelectron acceptor [oxygen)
is available eaergy geaeraticn frcm cx1ation cf substrate ac |cnger is pcssible an1 the
"bic1egra1ab|e" fracticn cf the ass |css a11s tc the eneshe1 5BC0D Tc supply aintenance
energy requireeats ua1er these ccn1i ticas pc|yP is c|eave1 [prccess 2l) Bcth pclyP cleavage an1
en1cgeacus ass |css are c1e||e1 as rst cr1er with respect to the pc|yP crgaaismass. Prcm the
eahaace1 cu| ture kinetic c1el, fcr bcth aercbic aa1 aaaercbic]ancxic states , the po|yP an1 PBB
ccnteat cfthe active asslcstisrelease1tc soluticn, thepclyP a11iag tcthe scluble P pcol an1the
PBB tceaeshe1C0D[prccesses 2I an122respectively)
Iathe NDBPRkiaetic c1e| , interacticns between the crgaaism pcpu|aticns are acccc1ate1thrcugh
the stcichicetric re|aticnships between prccesses an1 ccpcua1s - ac 1irect interaction processes have
been i1eatie1 Tc acccc1ate the interacticas betweea pc|yP an1 acn-pc|yP organiss the
stcichicetric re|aticnships between prccesses an1 ccpcun1s carrie1 cver frc the enhance1 cu|ture
kiaetic c1e| tequire1 re|ative|ymiacrchanges Intheeahance1 cu|ture kiaetic c1e| ,
[ I) the stcre1 PBB ccateat cf the pc|yP crgaais bic|cgical active ass 1isappeariag in aercbic aa1
aaaercbi c]aacxi c 1cgeacus mass |css was re|ease1 as acetate, it is ncw release1 tc eneshe1
5B00D,
[2) ua1er anaercbic an1 ancxic ccn1iticns, the bic1egra1ab|e pcrtica cf the pc|yP crganis bic|cgical
active ass 1isappeari ag ia ea1cgeacus ass lcss was treate1 as "uabic1egra1able", it is ncw a11e1
tcthe eaeshe1 5B00D,
[l) the mtrcgen ccntent cf the bic1egra1ab|e pcrticn cf the pclyP ctgaais bic|cgic active mass
1isappearing 1uriag aerobic an1 anaercbic]ancxc endcgeacus mass lcss was a11e1 to the ammonia,
itisacw a11e1 tcthe bic1egra1able particUate organic nitrcgen.
70
C.
60
.
50

o
: 40

a
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30
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20
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MI XBd CuI l ufB8
- b8CBd CuI l ufB8
20 30 40 50 60 70
MEASURED P REMOVAL (mgP/I )
PgI0. Pre1icte1 versus measure1 P recval, pre1icticns usiag NDBPR kinetic c1el, 1ata frcm
stea1y statelabcratcryce systems.
NBEPR systems
79
vuuatonoftheNDBEPR Hnetic model
AtpresenttheNDBPRkinetic c1elis beingevuate1, theevaluatonstll is restrcte1tostea1ystate
coa1itioas fcr reascn that extensive stea1y state experient 1ata are available (Wenttel et al. , I) .
This evaluation al|ows verication or o1ication ofconstaats carrie1 over frc the ND an1 enhance1
cu|ture kinetic ode|s. Iro thesiulaticns un1ertaken thus far, it appears that thevuesfcrconstants
fro the ND an1 enhance1 culture kinetic odels can be ret ained, with the exception of p_. An
iportant aspect cf the eva|uaticn has been to investigate the value for p_ over the wi1e range of
congurations an1 ccn1itions. The sulations for a particular syste are repeate1 usng a series of p_
va|ues unti| the experientally easure1 nitrateconcentrations are clcsely pre1icte1. Iro the set of p_
va|ues obtaine1 fcr the 1ifferent systes, the "best" p_ value is est| ate1. Iro the siulations
ccplete1 ( ! cf hich ! ccul1 be use1 to evaluate p_) , a ean value fcr p_ of. has been cbtaine1,
with stan1ar1 1eviaticn cf the ean + . l. 0sing the in1ivi1u p_ values that give the correct nitrate
concentraticns fcr aparticularsyste, thecorrelat|onbetweenpre1icted an1easure1P reovfor both
theixe1 an1enhance1cu|turesystesisshownin Iig I.
AspectsoftheNDBEPR Hneticmo1etobecomplete1
AnubercfaspectsoftheNDBPRkineticc1el requirefurther atteat| ca.
(1) 0op|eticn oftheo1e| evaluation un1er stea1y state ccn1i t|oas, i n part|cu|ar quanticaticn of p
(tota| 1atasets availab|e I I I ) .
(2) valuation of the o1e| un1er cyclic ow an1 loa1 con1i tions - a proble here is the lack of
coprehensive experiental 1ata for NDBPRsystes un1er cyclic ow an1loa1 (onlyonecoplete
cat aset is avai|able) .
(3) Denitricaticn by polyP crganiss - in the NDBPR kinetic o1el , |ro experiental observations
(Wentzel et al. , Ia, 0layton et al. , I, II ) 1enitr|catioa by pclyP organiss is not i nclu1e1.
Bowever, n soe instances sign|cant P uptake in the anoxic reactcrs ofNDBPRsystes has been
observe1, i p|ying that cn occasion the

c|yP organiss 1o 1eni tr|fy. Biochecal assays have


in1icate1 that soe Acinetobacter specesstrains have the ability to 1eni tr|fy ( Ltter, I) . At
present , no conclusive ju1geent can be a1e as to what con1iticns wll in1uce the presence of
1enitrifying polyP crganss in the NDBPRsyste. The oHy gui1e is om thermodynaics , that
1enitrication etabolis is energeticly less favourable than aerobic etabolis, and acccrdingly it
wou|1see that thepresenceof1enitrifyingpolyP organiss is anunstabletransiticnaryoccurrence.
[1) Teperature 1epen1ency- no quanttative inforationis availableon the teperature depen1ency of
the reactions e1iate1 by the pclyP organiss (e. g. aru specic grcwth rates, etc. ) . Bowever,
quali tativein1ications arethattheass ofP reovalis relatively teperaturei nsensitive.
() Behaviour ofnon-pclyP an1 autotrcphic crganiss un1er P liitation- i n the odel, following the
experiental observations an1 conclusicns ofWentzel et al. ( Ia, Ib) , shcul1soluble P becoe
liiting the po|yP organiss util|te polyP as an alternatve P sourceforcell growth. No nforaticn
is avai|ab|e as to the behaviour ofthe ncn-polyP an1 autotrophic organss un1er these con1tons -
inthe c1e| it is assue1 that growthcfthese organissceases un1er P liitingcon1i t|ons.
CLOSURE
Iro this review it woul1 appear that in the near future a relat|vely coplete ki netic description of
NDBPRactivate1slu1ge systes willbeavailable, a1equatefor,
i1enticatonofopt1esignproce1ures ,

estiation of tie response cf proposed design under dynac 0ow and load condtions, to assist in
1esigno1caton, an1
assessent ofcontrolstrateges tobe pleente1 at erstngorpropcse1 plants to opt|itenutrent
reoval.
0learly, BPRis now rly establshe1 and the echanss reasonably well un1erstoc1. Research ay
provide greater clarity on thephenomenabut technicly there is now a1equate understan1ing to exploit
PR at fullca|e, the future ofBEPR ao longer depen1s on better un1erstanding ofthe phenomenon,
but on better understan1ing of how to deal with the probles that can arise i n operation of NDBPR
plants. Theseprobles ainlyare.
80 M. C. WETL 6l al.
slu1ge bulking,
ina1equate RB00D in the in0uent tc give require1 BPR [ cne sclnticn, fcr example, is RB00D
generaticnbyaci1fermentaticncfprimaryslu1ge) ,
peric1s cf re1uce1 BPR efciency [cne scluticn is prcvisicn cf back-up systems such as chemical
a11iticn,
slu1ge han1ling an11ispcsal, an1
practical 1esignaspects tc explcit thisnew technclcgycptimally.
ach cf these prcblems i s being actively investigate1, but a review cf thei r present status wcU1 be
extensivean11cesnctliewithinthe sccpecftHspaper.
ACKNOWLEDGEMENTS
This research was suppcrted jcintly by the Water Research 0cmmi ssicn cf 5cuth Africa an1 the
Icun1aticnfcrResearchDevelcpment an1is publishe1 with theirpermissicn.
RFERNCES
Arvin, B [ I5) . Biclcgical remcv cfphcsphcrus frcm waste water. )0 0riticg Rev. Bnvircnmental
0cntrol , I5, 211.
Barnar1,J I [ II1) . 0utP an1 N withcut chemics . Water an1 Wsts Bnineerin, i, 33-3.
Barnar1, J I [ II5) . Nutrient remcvalinbiclcgic systems. Wat. PcUut . qntrol, I1[2) , I13-I51.
Barnar1, J I [ II). A review cf biclcgical phcsphcrus remcval in the activate1 slu1ge prccess.
W ter5 , [ 3) , I2-I11.
Bergey[ I1 . Hangcf1eterminativcteriolc_ [ I1thB1). Williams an1Wilkins 0c. , Baltimcre
Brc1isch, K B 0 [ I5). Interacticn cf 1ifferent grcups cf microrgasms in biclcgical phcsphate
remcval . Wat . 5ci . Tech. , II[ II]I2) , I3-I1.
Brc1isch, K 0 an1 Jcyner, 5 J [ I3) . The rcle cf mi crccrganisms cther than Acinetobacter in
biclcgicphcsphateremcval inactivate1slu1geprccesses. Wat . 5ci . Tech. , 1 (3/4), 1 1 7-1 25.
Buchan, I [ II). Thelccaticn an1 naturecfaccumulate1 phcsphcrus i nseven sIudges trcm actvate1
slu1geplants whichexhibite1enhance1phcsphcrus remcv. Water 5A, 1, I-I.
Buchan, I [ I3) Pcssiblebiclcgicalmechanismcfphcsphcrusremcval. Wat . 5ci . Tech. , 1 (3/4) , 87-1 03.
0laytcn, J A, kama, G A, Wenttel, H 0 an1 Harais, G v R [I). Demtricaticn kinetics in
biclcgicnitrcgen an1 phcsphcrus remcv activate1slu1ge systems . ResearchRe_crt W3, Dept .
0ivil ng , 0niv. cf0apeTcwn,Rn1ebcsch, II00, 5cuth Africa.
0laytcn, J A, kama, G A, Wenttel, H 0 an1 Hars, G v R [ II ) . Detricaticn kinetics in
biclcgical nitrcgen an1 phcsohcrus remcval activate1 slu1ge systems treating mucipal
wastewaters. Wat . 5ci . Tech. , n (4-6), 1 025-1 035.
0cmeau, Y, Ball, K J, Banccck R W an1 0l1ham, W K [ I). Bicchemical mc1el fcr enhance1
biclcgical phcsphcrus remcval. Proc. BC Conference cn new ethc1s an1 research in waste
treatment an1 resi1uals management , June, Vanccuver, 0ana1a Pnb. inWater Research, 2[ I2),
I5II-I52I I) .
Deinema, D, Baets, L B A, 5chclten, J, Turkstra, an1 Webers, H A A M [ I0) . The accumulaticn
cfpclyphcsphateinAcinetobacter spp. IH5Hicrcbicl . Ietters, j, 2I5-2I.
Dcl1, P I, kama, G A an1Harais, G v R [ I0) . A general mc1el fcr the activate1 slu1ge prccess.
Prc. Wat . Tech. , 1 2 (6), 47-77.
Dcl1, P I, Wenttel, H 0, Billing, A , Wenttel, H 0, kama, 0 A an1 Harais, G v R [ II).
Activate1 slu1e simulaticn _rcrams . Pub. by Water Research 0cmmi ssicn, P 0 Bcx 21,
Pretcria, 00I, 5cuthAfrica.
Dcwning, A I, Pnter, B A an1 Kncwles, G [ I1) . Nitricaticn in the activate1 slu1ge prccess.
J. Prcc. Inst . 5ew. Purif. , 1, I 30-I5.
kama, G A an1 Hars , G v R [ II) . Dynamic behavicur cf the activate1 slu1ge prccess.
J. Wat . Pcll. 0cntr. Ie1. , 5I, 31-.
kama, G A, vanBaan1el, A 0 an1Hars, G v R [ II). The present statuscfresearch cn nitrcgen
remcv. a mc1el fcr the mc1ie1 activate1 slu1ge prccess . 5ympcsium cn nutrient remcval, 5A
Branch cf IWP0, Pretcria, Hay. Prcc. gub. b_ Water Research 0cmmissicn, P 0 Bcx 21,
Pretcria, 000I, 5cuthAfrica.
kama, G A, 5iebritt, I P an1 Harais, G v R [ I3) . 0cnsi1eraticns i nthe prccess 1esigncf nutrient
remcv activate1 slu1geprccess . Wat. 5d. Tech. , 1(3/4), 283-3 1 8.
Plcrentt, H an1 Bartemann, P [ I2). Bntymatic stu1y cf activate1 slu1ge in aercbicnaercbic run.
Bnvircn. Tech. Ietters, , 315-350.
Puhs, G W an1 0hen, M [ II5). Hicrcbiclcgical buis cf phcsphate remcv in the activate1 slu1ge
prccessfcr the treatmentcfwastewater. Hicrcbicl. Bcclc@,, I I-I3.
Pukase, T, 5hibata, H an1 Hijay, X [ I2). 5tudes cn the mechani sm cf biclcgical phcsphcrus
remcv. Ja_an J. Water Pcllut. Rs. , Q, 30-3II[Trans. by0 J Har1cn, Newslettercf the stu1y
grcupcnphcsphateremcvinbiclcgicsewagetreatmentprccesses, 2 I ) , Peb. I1)I .
NBEPR systems
8 1
0erber, A, 1eVilliers, R B, Hcstert, S an1van Riet , 0 J J [ II) . The phencencn cfstultanecus
phcsphate uptake an1 release, an1 its ipcrtancein biclcic nutrient recval. Pub. |nA1vanes
in Water PclluticnQl ol: Bioloical Phosphate Removatfi"Oll1 Wastewaters, 1. R Raa1cri,
PergacnPress, 0xfcr1 pp. 1 23-1 34.
bl, , Nichclls, B A an1 0sbcrn, D W [ II) . Prcgress in Jchannesburg tcwar1s the biclcgic
recv cf phcsphcrus frcm sewage eMuents. I Ith ssen 0cnJ,, Inst cfWater Quali t an1 Water
ualitHanaeent .
bart, H A an1 Hele1, I N I2). Hicrcbiclcgy cf nutrient recving activate1 slu1ge Wat . Sc|
Tech. , 1 4 (9-1 1 ), 1 501 -1 502.
basccet , H 0, Ilcrentt, H an1 Cranger, P [ I5 . Biccheical aspects cf enhance1 biclcgical
phcsphcrus remcvfrcmwastewater. Wat . Sci .

ech. , II[ II]I2) , 2l~I .


bente, H, Cra1y, 0 P I [Jr), Cujer, W, Harai s, C v R an1 Hatsuc, T [ II). Activate1 Slu1geHc1el
Nc. I . IA WPR0 Scientican1 Technical Re_crt Nc. I , IA WPR0, Icn1cn.
lAWPR0 Task 0rcup [ II) . Task 0rcup cn "Hatheatical c1elling cf wastewater treatent " ,
meeting hel1i n0cpenhagen, August II.
1um [ II. 0eneticsan1physiclcgycfAcinetobacter. Ann. Rev. Hicrcbicl . , l2, l11-lII.
Iaw;ence, A an1 Hc0arty, P I [ II0) . 0nic1 basis fcr biclcgica| treatent 1esign an1 cperaticn
J. San. n. Div. , AS0, , I5I-II.
Ictter, I B [ I 5). The rle cfbacterial phcsphate metabclism in enhance1 phcsphcrus recv frc
theactivate1slu1ge prccess. Wat Sci . Tech. , II[ I I ] I2), I2I-Il.
.
Ictter, L B [ I). Dierl van
.
etabciesebeheer eanises in Acinet

bacter s__.

et betrekkmtct
fcsfaatverw_1erin 1eur 1re eaktiveer1esl_k_rcses. Dcctcral thest s, RA0 0mv. , Jchannesburg,
Scuth Africa. [inAfrikaans)
Ictter, I B an1 Dubery, I A [ I I). Hetabclic ccntrcl cf
P
hcsphate accu
.
mulatingbacteri

an1 its rle


in enhance1 biclcgical phcsphate recv. Pub. m A1vances m Water PclluttcnControl 4:
Biclcgical phcsphaterecvfrcmwastewaters, 1. R Raa1cri , PergacnPress, 0xcr1. pp. 7-1 4.
Ictter, I B an1 Dubery, I A [ I ). Hetabclic regulaticn cf hy1rcxybutyrate 1ehy1rcgenase in
Acinetobacter calcoaceticus var. lwoff. Water SA, I5[2) , 5-I0.

.
Marais, 0 v R an1 kama, C A [ II) . The activate1 slu1ge prccess. Part I - stea1y state behav:cur.
WaterSA, 2 I1-200.
Marai s, 0 v R, Icewenth, R an1Siebritt, I [ Il) . Review. 0bservaticns suppcrting phcsphate
recvbybiclcgical excess uptake. Wat . Sct . Tech. , I5[l]1) , I5-1I .
Massi11a, D, Wenttel , H 0, kama 0 A an1 Harai s, 0 v R II) . An investigaticn intc the
utilitaticn cf slcwly bic1egra1able crganic material in activate s|u1ge systes. Internal Re_crt ,
Dept. cf0ivlng. , 0niv. cf0ape Tcwn,Rcn1ebcsch, II00, Scuth Africa.
Matsuc, Y [ I5) . Iuncticning cf t he T0A c

cle un1er anaercbic ccn1i ticns in the anaercbic aercbic


acclimatite1 activate1 slu1ge [in Japanese. Prcc. cf10th Annual 0ca! cfthe Ja_an Scc. cf0iv.
@. , 1[2) , -0.
McKinney, R [ I2). Hathematics cfccmplete mixing activate1 s|u1ge J San. n. Div. , AS0, ,
I-I Il.
McKinney, R an1 0cten, R J [ I) . 0cncepts cfccplete mixing activate1 slu1ge. Trans. Ith
n. 0cnf. , 0niv. cfKansas , l2.
Meganck, H, Hncu, D, IeIlchic, P, Iaup, C H an1Rcvel, J H [ I5) . Theipcrtancecfaci1cgenc
icrc0crainbiclcgicphcsphcrusrecval . Wat . Sci. Tech. , II[ I I ] I2), I-2I2.
Minc, T, Arun V, Tsutuki , Y an1 Hatsuc, T [ II). ffect cf phcsphcrus accumulaticn cn acetate
metabcli s in the biclcgical phcsphcrus recval prccess. Pub. in A1vances in Water Pclluticn
Conrl 4: Biclcical phcsphate remcval frcm wastewaters, 1. R Raa1cri , Pergamcn Press,
0xfcr1. pp. 27-38
Murphy, H an1 Ictter, I B [ I). The effect cfacetate cn pclyphcsphate fcraticn an1 1egra1aticn
in activate1 slu1ge with parti cular reference tc Acinetobacter calcoaceticus: A icrcsccpic stu1y.
Water SA I2[2), l.
bichclls, B A (II5) . Iull scale experimentaticn atthenew Jchannesburg aeraticn plants. Water SA,
[l) , I2I-Il2.
bichclls , B A an1 0sbcrn, D W [II) . Bacteri stress. Prerequisite fcr biclcgical remcvu cf
phcsphcrus . J. Wat . Pcll. 0cntr. Ie1. , 5I[ l) , 55I-5.
bichclls, B A, 0sbcrn, D W an1 Pitman, A R [ II). Imprcvement tc the stability cfthe biclcgic
phcsphate remcv prccess at the Jchannesburg Ncrthern Wcrks. Puh in A1vances in Water
PclluticnCnt|4: Biclcgical phcsphate remcv frcm wastewaters, 1. R Rama1cri , Pergamcn
Press, 0xfcr1. pp. 26 1 -272
0sbcrn,D W, Ictter , I B, Pitman, A R an1 Nichclls , B A [ I). Twcyearstu1ycn theenhancement
cfbiclcgical phcsphateremcval by teringprccessf1 ccmpcsiticn [ Plant an1 labcratcry stu1ies) .
R_crt tc the Water Research 0cmmissicn, WR0 IlI/2/, P 0 Bcx 21, Pretcria, 000I , Scuth
Africa.
Rabincwitt, B an1 Hars, C v R [ I0). 0hemic an1 biclcgic phcsphcrus remcvalinthe activate1
slu1geprccess. ResearchRe_crt Wl2, Dept . cf0iv. ng. , 0nv. cf0apeTcwn, Rn1ebcsch, II00,
Scuth Africa.
82
M. C. WENTZEL 6l al.
Rensink, J B [ II) . Biclcgische 1efcsfatering en prccesbepalen1e factcren. In 5_. cek -
Defcsfaterin, nieuwe cntwikke|i nen en _rakti [kervarinen in Nederlan1 en 2we1en. NVA 5yp.
[inDutch) .
5iebritt, I P, kama, 0 A an1 Harai s, 0 v R I) . xcess bic|cgical phcsphcrus remcv in the
activate1 slu1ge prccess at warm temperate cliates . Prcc. Wasfe Treatent and 0tilitaticn, Vcl.
, 2ll-2I , 1. RcbinscnD W, Hcc-Ycung H an1Iarquhar0 J, Pergacn Press, Tcrcntc.
5iebritt, I P, kama, 0 A an1 Harai s, 0 v R Il). A paraetric c1el fcr bic|cgical excess
phcsphorus remov. Wat . 5ci . Tech. , 1 (3/4), 1 27-1 52.
9tern, I B an1 Harai s, 0 v R [ II1) 5ewage as the electrcn 1cncr in biclcgical 1enitricaticn.
Research Re_crt WI, Dept. cf0iBn. , 0niv. cf0apeTown, Rcn1ebcsch, II, 5cuthAfrica.
van Baan1el , A 0 an1 Harai s, 0 v R II) . Nitricaticn an1 1eni tricaticn ki netics intheactivate1
slu1geprccess . ResearchRe_crt l, ept . cf 0ivi| ng. , 0niv. c|0apeTcwn, Rcn1ebcsch, II,
5cuthAfrica.
vanBaan1el, A 0, kama, 0 A an1 Harai s, 0 v R [ II ) . Theactivatcd s|u1geprccess Part l - single
s|u1ge 1enitricaticn. Water Research, I, IIl-I I2.
van Baan1el , A 0, Dc|1, P L an1Harai s, 0 v R [ I2) . 0tiitaticn cfnitrcgen remcval inthesing|e
s|u1ge activate1 slu1geprccess. Wat . 5ci . Tech. , 1 (617), 443-46 1 .
Venter, 5 N , Lctter, L B , 1e Baas , D W an1 HacBcn1, L [ I) . The use cf the anytic prcle
in1exin the i 1enticaticn cf activate1 slu1ge bacteria. Prcbles an1 sc|uticns. Water 5A, I[1),
2-2I.
Warner, A P 0, kaa, 0 A an1 Harais, 0 v R [ I ). The act|vate1 sln1ge prccess Part 1 -
Applicaticn cf the gener kinetic mcdel tc ancxic-aercbic 1igesticn cf waste activate1 slu1ge.
WaterResearch, 2[ ). 1l-.
Watanabe, A, Hiya, A an1 Hatsuc, Y [ I1) . Labcratcry scale stadycn bic|cgi cal phcsphate remcval
using syntheti c wastewater. Recva| perfcrmance an1 the investigaticn cf enhance1 phophcrus
accumUating crganiss. Newsletter cfthe stu1 rcu_cn _hcs_hate recv inbiclcical sewae
treatment_rccesses, [ I ) , Iebruary.
Wentte|, H 0, Dcl1, P L, kama, 0 A an1 Harai s, 0 v R [ I). Kinetics cfbiclcgica| phcsphcrus
release. Wat . 5ci . Tech. , II, [ II]I2) , I-II.
Wenttel, H 0, Lctter, I B, Lcewenthal, R an1 Harai s, 0 v R [ I ). Hetabclic behavicur cf
Acinetobacter spp. in enhance1 biclcgic phcsphcrus remcv - a biccheical c1e| . Water 5A,
I2|1), 2-221.
Wenttel, H 0, Lcewenthal , R , kama, 0 A an1 Hars , 0 v R [ Ia). nhance1 pclyphcsphate
or

nism culturesin activate1 slu1ge systems- Part I. nhance1 cultnre1evelcpment. Water 5A


) , I-2.

Wenttel, H 0, Bkama, 0
.
A, D

l1, P I, Ioewenthal , R B an1 Harais, 0 v R [ Ib) . Biologic excess


phosphcrus remcv :n act:vate1 slu1ge systes. Research Re_ctt W, Dept. 0ivil ng. , 0mv. cf
0apeTcwn, Ron1ebcsch, II, 5outh Africa.
Wenttel, H 0, Dol1, P
.
I, Ioewenth, R , kama, 0 A an1 Harai s, 0 v R [ Ia) . nhance1
polyphcsphate orgamsm cultures :n actrvate1 slu1ge systes. Part II. xperiental behavicur.
Water5A, I[ 2) , II-.
Wenttel, H 0, Dcl1, P L, kaa, 0 A, an1Harais 0 v R [ Ib . nhance1 clyphcsphateorgamsm
culturesinactivate1s|u1gesystes . PartIII. Kineticmc1el. ater 5A, I 2) , -I2.
Wenttel, H 0, kama, 0 A, Dcl1, P Lan1Harai s, 0 v R [ I) . B|c|cgi ca| excess phcsphcrus reoval
- 9tea1ystateprccess 1esign. Water 5A, Jan. I.
Wentt

l , H , Lctter, I B, k

a, 0 A, Lcewenthal, R an1Harai s, 0 v R [ II) . Bvaluaticn cf


bicchemc mc1elsfor b:clcgical excess phosphcrusreov. Wat 5ci . Tech. , n (4-6), 567-576.
WR0 [ I1) . Thecr_, Desin an1 0_eration cf Nntrient Reova| Activate1 5|nde Prccesses. Water
Research 0ommission, P 0 Box21, 0I, Pretcria, 5cuth Africa.

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