Medicinl nd Aromtic Plnt Science nd Biotechnology 008 Globl Science Books Grlic Sprouts Grown Indoors t Kitchen Sites FeiFei Qin,,3* HuiLin Xu Gng M 4 Interntionl Nture Frming Reserch Center, Ht, Ngno, 390-40, Jpn Grdute School of Agriculturl nd Life Sciences, the University of Tokyo, Jpn 3 Penut Reserch Institute, Shndong Acdemy of Agriculturl Sciences, Qingdo, Chin 4 The United Grdute School of Agriculturl Science, Gifu University (Shizuok University), Gifu, 50-93, Jpn Corresponding uthor: * jiling_300@63.com ABSTRACT Recently, grlic shoots hve been used s vegetble nd functionl food in dditionl to the bulbs. In the present reserch, grlic sprouts were produced from grlic cloves with different sizes. The cloves were grown onto pper towels spryed with tp wter every two dys t kitchen sites under room temperture (fluctuted from 6C in the night to 6C round middy) or in growth chmber (5±/0±C, dy/night). The finl fresh nd dry biomss indicted tht the tiny cloves could lso produce edible sprouts though they were not s strong s those produced by lrge nd medium cloves. Fresh biomss ws mesured ech dy until hrvest nd the dt were used for nlysis of the dynmic growth of these grlic sprouts by sigmoid model, g G = g T {+(-t)e [-(t-)] }^(-)+g 0 (-t). One incidence nmed s Biomss Downcst Phenomenon occurred s shoot fresh biomss shrply declined when the nutrients in the clove were used up t the lter growth stges of grlic sprouts. The curve phse of the shrp declining ws nlyzed using n exponentil model, g D = g T +(g mx - g T )e [-(t-4)], nd the biomss downcst ws quntified using definite integrl formul, g () = *. Lef photosynthesis ws low due to the wek light in kitchen site conditions. It quickly reched light sturtion point (LSP) s the photosynthetic phonon flux (PPF) incresed to 50 mol m - s -. Photoinhibition occurred once PPF suddenly incresed bove LSP nd it ws quntified by definite integrl formul, P (i) = *. In conclusion, grlic cloves, even in smll size of no mrketing vlue, cn be used to produce fresh sprouts with high edible vlue t kitchen sites. The sprouts should be hrvested before the nutrition in the cloves is used up when the so-clled Biomss Downcst Phenomenon occurs. The mthemticl pproch dopted in this reserch for grlic sprout growth is of high reference vlue to plnt scientists. b ' ( gg() t gd() t ) dt ( PN( i) PI( i)) di * * b ' Keywords: Allium stivum, biomss downcst, growth, mthemticl pproch, modeling, photoinhibition, photosynthesis INTRODUCTION Sprouts from seeds of mny species re used s vegetbles becuse mny vluble functionl nd nutritive components re developed during germintion nd sprouting (Prk et l. 998; Tky et l. 003; Mizuno nd Ymd 006). Originlly, these sprouts were produced indoors with home sprouting kit t kitchen sites for fmily vegetble supplies. Sooner or lter, mss production systems were developed in response to the mrket demnd. Recently, concerns on helth nd food sfety increse nd led to dmirtions to sprout vegetbles s functionl food (Berry et l. 00). As consequence, species used in producing sprout vegetbles hve incresed from soyben (Prk et l. 998; Kylen et l. 007), mungben (Funki et l. 97; Kylen et l. 007; Kvs nd Sedef 008), green onion (Kuwy nd Whb 984), wtercress (Moriym nd Ob 004) nd rdish (Tky et l. 003) to lflf (Bhthen nd Velsquez 00; Kylen et l. 007), sunflower (Cho et l. 007), lentil (Urbn et l. 995; Kylen et l. 007; Kvs nd Sedef 008), pe (Urbn et l. 005), buckwhet (Kim et l. 004) nd grlic (Prk et l. 998). The merits nd dvntges in indoors production of sprouting vegetbles include the opportunity to try severl vrieties tht would enrich dinner tbles without being limited to sesonl vegetbles, efficient use of the seeds tht hve lost their economic vlue on the mrket, development of model for business, orgnic methods esily performed, nd the simple equipment of only couple of trys or kits insted of grden. Using sprouts is nother wy of mking vegetbles vilble in the off-seson nd expected vegetbles cn be produced to compenste the sesonlity nd fluctutions in supplies of the sme species. As the dded dvntges, the processing is home bsed, nd the seeds cn be stored nd sprouted esily during periods when vegetbles re scrce. As functionl food, grlic hs the highest ntioxidnt ctivity mong the common vegetbles nd sprouts bsed on the fresh weight (Co et l. 996). Grlic hs been used s both food nd medicine in mny cultures since the old ges. Grlic is climed to offer protection from cncers (Gleone et l. 006) nd prevention of hert disese (Ackermnn et l. 00) including therosclerosis (Berthold nd Sudhop 998), high cholesterol (Berthold et l. 998) nd high blood pressure (Pedrz-Chverrí et l. 998). In ddition to the medicinl nd functionl benefits, the tender sprouts from grlic cloves serve s fresh culinry nd sld, especilly the fresh ones mde indoors t the kitchen sites. In most cses, germintion nd sprouting need n pproprite wrm temperture to be mintined. However, with the exception, grlic dpts to brod rnge of temperture especilly t lower tempertures since it is winter crop in most res. Nevertheless, becuse of the lrger size, cloves of grlic contin lrger mount of nutrition nd re esier to generte strong sprouts compred to species with smll seeds. Grlic is one of kitchen grdener s drem crops, s it is plnted fter other crops re clered out in winter. Cold wether stimultes grlic cloves to root nd sprout (Nei et l. 007). It will quickly develop strong-tsting sprouts, nd Received: 9 September, 008. Accepted: 7 October, 008. Originl Reserch Pper
Medicinl nd Aromtic Plnt Science nd Biotechnology (), 7-008 Globl Science Books cloves cn supply enough nutrients for the young sprouts to grow (Mteljn 006). The tiny cloves mong the norml ones re usully not used for plnting nd often thrown wy s they will not mount to much other thn green grlic. However, tiny cloves might be preferred in kitchen site grdening. If the relly miniscule cloves cnnot be plnted in the field, they cn be plnted in moist continer in row firly close together, nd tender fresh sprouts cn be hrvested for use s spice nd sld culinry or for stirfries. Nevertheless, one problem is tht cloves grown in kitchen site conditions without soil nd enough light often produce wek nd yellow green sprouts or seedlings, which ffects the edible vlues. Considering the bove problem, in the present experiment, grlic cloves with different sizes were chosen s seeds for sprouting to mke sure whether the miniscule cloves cn produce norml sprouting greens in comprison with lrge nd medium cloves under kitchen site conditions. Mthemticl models were dopted to nlyze the growth dynmics nd photosynthetic ctivities of grlic sprouts. MATERIALS AND METHODS Grdes of grlic cloves Individul grlic cloves (Allium stivum L. Jinxing No. ) were seprted from bulbs nd sorted into seven grdes by size indicted s fresh biomss per clove. The seven grdes were s follows: Grde, 8.7 g; Grde, 7.04 g; Grde 3, 5.59 g; Grde 4, 4.8 g; Grde 5, 3.79 g; Grde 6,.70 g; nd Grde 7,.57 g. Cloves in seven grdes were further defined into lrge (Grdes nd ), medium (Grdes 3 nd 4) nd smll (Grdes 5, 6 nd 7) cloves on the size bsis. Tretments nd plnting Polystyrene pltes used in this experiment were 44 cm long, 3.5 cm wide nd 7.5 cm high. A lyer of white towel, seven lyers of guze nd 70 pieces of tissue ppers were pved t the bottom of ech plte s they could quickly bsorb wter nd mintin moisture for grlic sprouting. Ech plte ws divided into two prts tht were served s tretment repets, nd three pltes repeted for ech tretment. Cloves with the sme grde were plced closely in one row (Fig. ) nd the pltes were left open. The number of cloves in ech row ws different becuse of the different sizes in ech grde. In Grde, 0 cloves were grown per row; Grde, cloves per row; Grde 3, cloves per row; Grde 4, 3 cloves per row; Grde 5, 4 cloves per row; Grde 6, 7 cloves per row; nd Grde 7, 0 cloves per row. Tp wter of 500 ml ws spryed every two dys to ech plte to ensure the wter supply to grlic sprouts. Two temperture regimes were dopted in room (fluctuted from 6C in the night to 6C round middy) nd chmber (5±/0±C, dy/night) conditions, respectively. There ws no dditive illumintion supplement. The lighting condition in both room nd chmber ws 0-5 mol m - s - s the nturl light chnged during dy nd night. Anlyses of sprout growth nd biomss dynmics Five grlic sprouts of ech grde were smpled everydy until Dy 8 for mesurements of the fresh biomss. The finl fresh nd dry msses of seven grdes were recorded on hrvest dy (Dy 8 in chmber conditions nd Dy 8 in room conditions). As the whole fresh grlic sprout cn be used edible, dt of the totl fresh biomss were used for the nlysis of growth. A mthemticl pproch ws dopted to the nlysis of biomss ccumultion of during sprouting nd growing of the grlic sprouts. Mesurement nd nlysis of photosynthesis Photosynthesis ws mesured for nd compred between sprouts from the lrge nd smll cloves. Photosynthesis in the fully expnded leves of the grlic sprouts ws mesured using Li-6400 Portble Photosynthesis System (LI-COR Inc. Lincoln, Nebrsk, USA) on hrvest dy. Mthemtic pproches were dopted to nlyses of the photosynthesis nd photoinhibition in response to incresing light intensity. RESULTS AND DISCUSSION Fig. Grlic cloves grown in pltes t kitchen sites (tken on plnting dy). Sprout growth nd biomss dynmics The finl increment in fresh nd dry biomss of grlic sprouts indicted tht nutrients stored in grlic cloves could support the growth of cloves into edible sprouts when there ws only wter supplied nd no other substrte or nutrient solution dded. Although the finl increment in fresh nd dry msses in the sme grde showed no significnt difference between the room nd the chmber conditions (Tble ), high temperture dvnced grlic sprouts to rech edible stte t mximum biomss 0 dys erlier. In the present study, low temperture in room conditions delyed the growth nd it cost 8 dys to rech the mximum biomss in edible sttus. The finl biomss ws significnt different mong seven grdes. It ws fr less in smll cloves thn in lrger ones (Tble ). However, most importntly, even the tiny cloves of Grde 7 could lso produce sprouts in good shpe though they were not s strong s those produced by lrge nd medium cloves. Tble The finl fresh nd dry biomss (g sprout - ) of grlic sprouts under room nd chmber conditions. Grde Room (Dy 8) Chmber (Dy 8) Finl increment Finl increment Finl FM Fresh Dry Finl FM Fresh Dry.3 A 4.05 A 0.76 A.73 A 4.06 A 0.76 A 8.73 bb 3.0 AB 0.48 bb 9.8 bb 3.77 AB 0.60 bb 3 9.0 bb 3.69 bb 0.56 bb 8.54 cc 3.3 bb 0.49 cc 4 6.60 cc.9 cc 0.33 cc 6.46 dd.39 cc 0.4 dcd 5 5.5 dd.97 cc 0.33 cc 5.56 ee.39 cc 0.39 dd 6 3.77 ee.5 ccd 0.5 ccd 3.4 ff.5 dd 0.9 ee 7.3 ff 0.87 dd 0.4 dd.4 gg 0.68 ed 0. fe Condition ns ns ns ns ns ns Grde ** ** ** ** ** ** Condition Grde * * * * * * *, P0.05; **, P0.0; nd ns, no significnt difference ccording to ANVOA. Different letters show significnt differences t P0.05 nd P0.0 ccording to LSD test. 8
Modeling kitchen culture of grlic. Qin et l. Tble Prmeters of the sigmoid growth curves of grlic sprouts from cloves in different sizes under cold room nd wrm chmber conditions. Condition Grde g T (g) ( 0-3 dy - ) (dy - ) (dy) g 0 (g) Room 4.76 ± 0.6-0. ±.7 0.44 ± 0.06 8.88 ± 0.4 7.43 ± 0.05 3.67 ± 0.3-8.8 ±.87 0.57 ± 0.09 8.76 ± 0.4 5.8 ± 0.06 3 3.04 ± 0.4 -.84 ±.76 0.47 ± 0.04 8.48 ± 0.7 5.0 ± 0.06 4.70 ± 0.3.77 ±.74 0.36 ± 0.04 8.6 ± 0.5 3.8 ± 0.08 5.3 ± 0.6-0.03 ±.64 0.38 ± 0.05 7.65 ± 0.6 3.8 ± 0.08 6.39 ± 0.55 38.65 ± 6.78 0.3 ± 0.05 0.36 ± 0.79.43 ± 0.0 7 0.58 ± 0.09-5.08 ±.8 0.4 ± 0.08 8.73 ± 0.3.3 ± 0.3 Chmber 5.0 ± 0.75 0.67 ± 4.06 0.45 ± 0.07 9.96 ± 0.4 7.63 ± 0.0 4.6 ± 0.8.84 ± 4.99 0.34 ± 0.07 9.8 ± 0.3 6.7 ± 0. 3 3.6 ± 0.60.37 ± 4.4 0.43 ± 0.07 9.70 ± 0.8 5.5 ± 0.08 4 3.7 ± 0.8 4.9 ± 7.07 0.33 ± 0.08 8. ± 0.50 3.88 ± 0. 5 3.83 ±.44.47 ± 79.63 0.3 ± 0.08 8.4 ± 0.84.98 ± 0.45 6.40 ±.4 3.84 ± 9.59 0.6 ± 0.3 7.48 ±.0.8 ± 0.4 7.30 ± 0.66 3.3 ± 3.09 0. ± 0.09 5.83 ±.05.76 ± 0.48 Dynmic of sprout growth nd biomss downcst The dynmic chnges of the fresh biomss during the sprout growth were nlyzed using sigmoid model s follows: g G = g T {+(-t)e [-(t-)] }^(-)+g 0 (-t). Here, g T is the totl increment in fresh biomss by growth; g 0 is the originl fresh biomss of the clove; is constnt ssocited with other fctors thn the growth; is constnt ssocited with the fst growth; is the time point for the biomss increse to rech the hlf mximum; nd t is the growth time (dy) (Xu et l. 000). The fresh biomss of sprouts declined shrply t lter growth stge s the nutrients stored in cloves were used up. This incidence ws nmed s Biomss Downcst Phenomenon. The curve phse of the shrp declining ws nlyzed by n exponentil model s follows: g D = g T +(g mx -g T )e [- (t-4)]. Here, g T is the totl fresh biomss decrement in the biomss downcst; g mx is the mximum fresh biomss before it begins to decline; is the time constnt; nd t is the growth time (dy). The biomss downcst ws quntified using definite integrl formul: b ( g ( t) g ( t)) dt G D g () =, b where g () is the verge fresh biomss decrement due to biomss downcst; integrl intervl from to b is from 4 to 6; g G (t) is the sigmoid growth curve nd g D (t) is the exponentil decline curve; nd the right side of the eqution is the difference (-integrl re) between g G (t) nd g D (t). Although the growth curves showed some distinction mong the seven grdes, grlic spouts of ech grde grew in mnner of exponentil dynmic (Fig. ). The rpid growth strted minly on Dy 8 or Dy 9 (shown by the prmeter ). is proportionl to the fst growth of the sprouts. The bigger the vlue, the fster the growth. The vlue of in smll cloves ws much lower thn tht in lrge nd medium cloves (Tble ), which indicted the slower growth in smller cloves. Fresh biomss ccumultion of grlic sprouts under room conditions declined nd even shrply decresed t the lter growth stges. This shrp decrese in fresh biomss ws more obvious under chmber condition with the higher temperture regime (5±/0±C, dy/night) (Fig. 3). The sudden decrese in fresh biomss, the phenomenon clled biomss downcst, begn on dy 4 when the fresh biomss reched the mximum, nd before tht time the growth followed the sigmoid model. Although higher temperture promoted the growth of grlic sprouts nd shortened the time for sprouts to rech Fresh Biomss per Sprout (g) 3 0 9 8 7 6 5 4 3 0 5 0 5 0 5 30 Dys of Growth Fig. Sigmoid growth curves for grlic sprouts from cloves in seven different sizes under cold room condition. From the top down: Grdes,, 3, 4, 5, 6 nd 7. Fresh Biomss per Sprout (g) 3 0 9 8 7 6 5 4 3 0 5 0 5 0 5 30 Dys of Growth Fig. 3 Sigmoid growth curves for grlic sprouts from cloves in seven different sizes under wrm chmber conditions. From the top down: Grdes,, 3, 4, 5, 6 nd 7. The lst four dt (on dy 6, 8, nd 6) in the upper prt of ech curve were nlog dt ccording to the sigmoid growth curves from dy 0 to dy 4, nd the four dt in the lower prt of ech curve were the rel dt tht followed exponentil declining curves. hrvestble sttus, nutrients stored in cloves were depleted quickly s the growth of grlic sprouts progressed. Furthermore, photosynthte t the low light conditions ws hrd to supplement the depletion by respirtion nd it ws the high respirtion tht resulted in n mzingly fst growth t the middle stge (Fig. 4). High temperture ccelerted respir- 9
Medicinl nd Aromtic Plnt Science nd Biotechnology (), 7-008 Globl Science Books Tble 3 Prmeters of the growth declining curves of grlic sprouts under chmber condition from dy 4 to dy 6. Grde g T (g) g mx (g) (dy - ) Biomss downcst * g () (g) 0.64 ± 0.3.0 ± 0.8 0.65 ± 0.9 8.99.58 8.66 ± 0.5 9.79 ± 0.3 0.84 ± 0.65 4.9.4 3 6.85 ± 0.50 8.75 ± 0.5 0.5 ± 0.08 3.75.5 4 5.7 ± 0.0 0.58 ± 0.4 0.60 ± 0.30 0.5 0.88 5 4.5 ± 0.03 5.74 ± 0.04 0.37 ± 0.04.90.07 6.64 ± 0.04 3.83 ± 0.05 0.49 ± 0.06 3.60.34 7.8 ± 0.03. ± 0.04 0.47 ± 0.5 4.70 0.39 Tble 4 Photosynthetic cpcity (P C), quntum yield (Y Q) nd drk respirtion rte (R D) of grlic sprouts under different environment conditions. Condition Size P C (mol m - s - ) R D (mol m - s - ) Y Q (mol mol - ) Room Lrge 6.39 0.6 0.05 Smll 5.76.3 0.030 Chmber Lrge 5.0 ± 0.43 3.9 ± 0.36 0.03 ± 0.000 Smll 5.3 ± 0.66 4.55 ± 0.56 0.07 ± 0.00 Photosynthesis( mol m - s - ) tory depletion during the lter growth periods (Fig. 5). Tht ws why the biomss downcst occurred to lrger extent in the chmber conditions. Lrger cloves produced thicker nd stronger sprouts in which the respirtion (respirtion rte lef re) ws higher thn tht in thinner nd weker sprouts. The vlues of biomss downcst * nd the verge fresh biomss decrement (g () ) were high in lrge nd medium grlic sprouts, nd only 4.70 in biomss downcst nd 0.39 in g () in the sprouts produced by miniscule cloves (Grde 7) (Tble 3). Biomss downcst ws the min reson for nutrition loss of grlic sprouts. Therefore, growing grlic sprouts in kitchen site conditions should void the occurrence of biomss downcst nd grlic sprouts tht hve reched edible sttus should be hrvested in time before the biomss downcst occurs. b 0 - - -3-4 -5-3.9-4.63 * ( gg( t) gd( t)) dt Fig. 4 grlic sprouts grown under cold room (left) nd chmber (right) conditions t lter growth stge (tken on dy 6). Lrge Smll 0 500 000 500 000 Photosynthetic photon flux( mol m - s - ) Fig. 5 The response curves of photosynthetic rte to the incresing PPF for grlic sprouts grown under wrm chmber conditions. The intersection points tht the curves crossed with Y-xis were respirtion rte (R D). Photosynthesis nd photoinhibition The response curves of photosynthetic rte to the incresing photosynthetic photon flux (PPF) were nlyzed using n exponentil eqution: P N = P C (-e -Ki )-R D. Here, P N is the net photosynthetic rte; P C is the mximum gross photosynthetic cpcity; R D is drk respirtion rte; K is constnt, equivlent to the reciprocl of the PPF t which P N reches 63% of P C ; nd i is the PPF nd the mximum quntum yield ws shown s Y Q = KP C (Togri 973; Steven 998; Xu 000). Photosynthetic rte in leves of grlic sprouts ws low s result of the wek light in room conditions (Figs. 5, 6; Tble 4). Another photosynthetic phenomenon clled photoinhibition ws observed in grlic sprouts grown under room conditions in the present study. The curve phse of the fst declining by photoinhibition ws nlyzed using n exponentil formul: P I = P 0 +P D e (i-50). Here, in lrge cloves P 0 is the mximum photosynthetic rte before photosynthesis begins to decline, nd in smll cloves P 0 is the minimum photosynthetic rte fter photoinhibition; P D is the decresed photosynthesis nd is constnt (Xu et l. 000). The photoinhibition ws quntified using definite integrl formul: P (i) = ( PN( i) PI( i)) di ', ' where P N (i) is the photosynthetic light response curve tht would be without photoinhibition nd P I (i) is the curve of photoinhibition; integrl intervl from to b ws from 50 to 000 mol m - s - ; ' ( P ( i) P( )) N I i di is the difference (-integrl re) between P N (i) nd P I (i). Photosynthetic cpcities of grlic sprouts were low becuse there ws lck of enough light in both the room nd chmber conditions. Photosynthesis in the sprouts grown under room conditions quickly reched the light sturtion point s the phonon flux incresed to 50 mol m - s - (Fig. 6). Photoinhibition, the reduction in plnt s cpcity for photosynthesis, is often cused by exposure to strong light tht is bove the sturtion point (Nom et l. 003). In the present study, low lighting, deficiency of nutrients nd the low temperture under room conditions together resulted in the serious photoinhibition phenomenon once the PPF suddenly incresed to level bove 50 mol m - s -. In ddition, photoinhibition in smll cloves ws greter thn tht 0
Modeling kitchen culture of grlic. Qin et l. Tble 5 Prmeters nd definite integrl showing the chrcteristics of photosynthetic declining cused by photoinhibition in grlic sprouts under cold room conditions. Size P 0 P D P (i) (mol - m s) ( ( ) ( )) mol m - s - ' N I P i P i di Lrge 3.37-0.09.9 3349.76 0.003 Smll 0.53.48.0 3847.99-0.0035 Photosynthesis( mol m - s - ) 6 5 4 3 0 - - -3-0.6 -.3 in lrge cloves (dt of P (i), Tble 5). The possible resons might be the high respirtion rte (.3 mol m - s - ) nd low clove nutrition supply in thin sprouts produced by smll cloves. Although the photosynthetic cpcity ws low in grlic sprouts grown under chmber conditions, photoinhibition did not exist becuse the sprouts dpted to the high temperture tht might be ssocited with high light (Fig. 5). CONCLUSIONS Lrge Smll 0 500 000 500 000 Photosynthetic photon flux( mol m - s - ) Fig. 6 The response curves of photosynthetic rte to the incresing PPF for grlic sprouts grown in cold room conditions. The dsh prts of the curves were modeling curves without photoinhibition. The solid lines show the rel photosynthetic declining cused by photoinhibition s PPF incresed. The intersection points tht the curves crossed with Y-xis were respirtion rte (R D). The experiment confirmed tht tiny cloves of grlic without seed nd mrketble vlues could be used to produce grlic sprouts in good shpe. Wrm tempertures by heting systems nd supplementl lighting were not necessry for grlic production nd the sprouting could be performed even under the cold room conditions. Sprouts should be hrvested before the stored nutrition in the cloves ws used up when the so-clled biomss downcst might occur. In the present study, mthemticl methods were used to model nd nlyze the dynmics of growth nd biomss ccumultion in the grlic sprouts. The equtions nd the nlyticl methodology re vluble in reference to plnt scientific reserch. One who grows grlic sprouts in the kitchen site conditions should be dvised s follows: ) grlic sprouts need light even in wek sttus in the room conditions; ) temperture should be not high in voidnce of excessive growth; nd 3) hrvest t optimum edible stge. In this experiment, Dy 6-8 ws the best hrvest time for grlic sprouts grown under cold room conditions, nd Dy 4 for sprouts grown in the chmber under 5±/0±C (dy/night) becuse of the biomss downcst. Although the illumintion ws low in room conditions nd consequently resulted in wek photosynthetic ctivity, the sprouts developed leves in good shpe nd norml green in the cold room conditions. At lest, light ws needed for the cholorophyll formtion in the sprout leves, no mtter how much the photosynthte contributed to the biomss in ddition to the contribution from the nutrition supplied by cloves s the min sources. 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Medicinl nd Aromtic Plnt Science nd Biotechnology (), 7-008 Globl Science Books List of symbols nd bbrevitions used in this study Symbol Unit Definition LSP Light sturtion point PPF mol m - s - Photosynthetic photon flux g G g Fresh biomss t given growth time g T g Totl increment in fresh biomss g 0 g Originl fresh biomss of the clove dy - Constnt ssocited with the fst growth Constnt ssocited with other fctors thn growth dy Time point for the biomss increment to rech the hlf mximum t dy Vrible of growth time g D g Biomss t given time during the biomss downcst g T g Totl fresh biomss decrement in the biomss downcst g mx g Mximum fresh biomss before the biomss downcst Constnt ssocited with biomss declining in biomss downcst g () g Quntity of the verge biomss decrement due to biomss downcst g G (t) g Sigmoid growth curve g D (t) g Exponentil biomss declining curve P N mol m - s - Net photosynthetic rte P C mol m - s - Mximum gross photosynthetic cpcity R D mol m - s - Drk respirtion rte K m s mol - Constnt for photosynthetic light response curve, equivlent to the reciprocl of the PPF t which P N reches 63% of P C i mol m - s - Vrible of photosynthetic photon flux Y Q mol mol - Mximum quntum yield shown s Y Q = KP C P I mol m - s - Photosynthetic rte t given PPF during photoinhibition P 0 mol m - s - Photosynthetic rte before or fter photosynthesis begins to decline due to photoinhibition P D mol m - s - Mximum decrement in photosynthesis due to photoinhibition mol - m s Constnt ssocited with photoinhibition P (i) mol m - s - Quntity of the verge photosynthesis decrement due to photoinhibition P N (i) Photosynthetic light response curve tht would be without photoinhibition P I (i) Curve of photoinhibition