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MAINTENANCE RESPIRATION in CROP LEGUMES

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MAINTENANCE RESPIRATION in CROP LEGUMES IVAITE INSTITUTE 13'lo 81k LIBRARY MAINTENANCE RESPIRATION iN CROP LEGUMES DONALD E. IRVING B.Sc., M.Sc. Hons. (Auckland) A Thesis submitted for the degree of Doctor of PhilosoPhY in the Faculty of Agricultural Science DePartment of Agronorny, l-laite Agricultural Reseàrch Institute UniversitY of Adelaide 1983 To my l'lother, my late Father, ny Sister and my Brother. CONTEN iS Page Nq. CHAPTER I GENERAL INTRODUCTION 1 CHAPTER II LITERATURE REVIE'.¡l 6 TNTRODUCTTON 6 PATHWAYS 0F C02 EV)LUTÏON 6 Pathuays of Heæose )æid'a.tion 6 qLucolusis 7 the oridatiue pen tose ohosphate pathuau 7 the Lic aeid Le 7 7ther Pathuays InuoLtsed in C0, EuoLution I MII)CH)NDRIAL ELECTR)N TRANSP1RT PATH\"-IIS 9 THE CONCI]PT OF GROI¡TH AND MATNTENANCE RESPIRATION 10 Hi stoz"ilaL Intro ductí on 10 Recent DeueLopments in the Concept of Grouth ortd MeLnteTltmce Respínation 11 Ihe WderLyíng Significance of the Grouth Effiaieney øn'd Maintenøt ce C o e f fi aL ent 74 qrouth effiaieneu t4 the maintenøtee eoeffieient 16 ErperimentaL Deternination of Gtouth øtd MqintenqrLee Coeffieients 16 Assurnptíons Made in CaLeuLating Cz,outh øtd. Maintencnce Coeffíeients 1B Faetov,s Affeetíng the Maintenqnce Coeffieient 20 Ineonsisteneies ín the Concept of Ì,lainterrcmce 2t Py,aeticaL AppLications of the Gv'owth and Mcrintenøtee Coneept in PLøtt Ssience 23 CONCLUSTON 24 CHAPTER III MATERIALS AND METHODS 26 PLANT CALTURE 26 WOLE PLANT GAS EXCHANGE 27 LEAF GNS EXCHANGE - INFA¿, RED GAS ANAT'YSIS 27 LEAE GAS EXCHANGE - BY MA]VOMETRY 28 PLANT ANALYSIS 28 Dzy Weight 28 I'eaf Area 28 ChLorophyLL 28 Carbon Content 2B Nítrogen Content 29 Protein Assøy 29 methods used in eæper"iment one 30 P_age No. soluble protein 30 insoluble protein 30 total protein 30 RUBPC 30 ri f i cat'ion 30 anti body rrroducti on 31 rocket i mmunoel ectrophores i s 31 metho,Ts used in eæpev"iment k¡o 32 Stareh Measurement 32 sønpLe pz,eparation 32 s tartdard p z' ep arati on 33 øruLoq Luco si das e pz'ep arati on 33 gLueose assalt 33 SoLthLe Carbolty drate Measu.renent 33 CHAPTER IV INTERRELATIONSHIPS BITWEEN RESPIRATION IN PROLONGED ÐARKNESS, DRY '/lEIGHT AND NITROGEN CONTENT 34 TNTRODUCTION 34 METHODS 34 RESULTS 35 WhoLe PLøtt Stu&ies 35 Root øtd Shoot Studíes 37 v.oot and shoot nespíra tíon in proLorwed døvkness 37 nitv,oqen coneentratíon in roots øtd shoots 37 DTSCUSSION 38 CONCLUSTONS 40 CHAPTER V APPLICATION OF THE CONCEPT OF GROI^ITH AND MAINTENANCE TO CROP PLANTS 47 LN?RODUCTTON 4t METHODS 4t CaLcuLation of Gt'outh ø¿d MainterrcInce Cceffiaients by the funønic Method 42 CalcuLation of Grouth and Mqintenance Coeffíeí'ents by the Steady State Method 43 CalcuLatíon of Grouth øtd l,laLntenØtee Coeffíe|ents by the Gtoss Uptal<e Method 44 RESULTS 44 Tne lulaintenance Ccefficí ent Íh.e Groubh øffieieney DTSCUSSION 45 The MaintenúIee Coeffíeient 45 The Grouth Effieieney 49 Effect of Species 49 CHAPTER VI DARK RESPIRATION OF MATURE LEAVES DURING PROLONGED DARKNESS 51 TNTRODUCTÏON 51 IúETHODS 51 Grout'!, Tv'e at¡ne-nts 51 Procedures 52 RESULTS 53 Ihe Relatíonship betueen Dark Respiration ø¿d Preuious Irz,adùqnce I'eueL 53 Leaf Respiration in Pt'oLonged Daz'kness 53 effeet of pv'euious ív'ra&Lanee 53 effeet of Leaf detachment 54 effect of debuddinq 56 effect of aLtez"Lnq the nitnoqen suppLu 57 the qas auotient 58 CaLeuLatíon of the Errcngy RequLrement fot' Suez'ose Irøtspoz't 58 CaLeuLation of the MaLntenØLce RequLrement fon Mature Ieaues 59 DTSCUSSTON 61 CHAPTER Vii CARBOHYDRATE EXPORT AND SOURCES OF SOLUBLE SUGARS FOR LEAF RESPIRATION DURING PROLONGED DARKNESS 67 TNTRODUCTÏON 67 METHODS 68 RESULTS 69 Ihe Time Coru,se of WhoLe PLøú Carbon Eæchange FoLLouíng DefoLiatíon 69 Ílte TLme Cowse of Carbohyd.nate Concentv,atíon in the Mature Leazses 70 The Carbohydrate PooL fon Leaf Respiration 72 DÏSCUSSION 72 þge lio. CHAPTER VIII VARIATION IN THE PP.OTEiN CONTENT OF MATURE LEAVES DURING PROLONGED DARKNESS 79 79 TNTRODUCTION METHOß 79 RESULTS BO Tlp TLme Course of Leaf ChLorophylL Content 80 TIp ?ime Cowse of Leaf Protein Concentratíon B1 solubLe protein 81 ínsoLubLe ptoteín 83 totaL proteín 85 Tl¿eoreticaL Protein Degtadntion Rates 87 Recouezn¿ of Photosynthesis Aftet' a ProLonged natk Treat¡nent 87 88 DTSCUSSÏON CHAPTER IX GENERAL DISCUSSION 92 TilETHODS FOR THE WASUEEMENT OF AAINTENANCE RESPIRATTON 92 MATURE LEAE RESPIRATTON AS REPRESENTATIW OF THE INTENSITY OF THE MLTNTENANCE PROCESSES 96 SUGAR POOLS FOR GROT^ITH AND MLTNTENANCE RESPIRATTON 98 PATH\IAY HWUTHESIS AND PROSPECTS FOR BREEDTNG lHE ALTERNATTW 99 , LOW, MATNTF:NANCE PLA],ITS APPENDIX I 101 RTFERINCES CITED L02 SUI4MARY Maintenance respiration refers to energy production for processes other than the synthesis of ¡rew ceìl nraterials, and it follows therefore that as the energy requirements of maintenance processes increase, the quantity of carbon allocated to growth is likely to be proportionately decreased. l,Jhilst the quantitative relationship between respiration (dark C0, efflux) and growth has been extensively studied, considerably fewer studíes have attempted to quantify the energy require- ments of the maintenance processes and experiments reported in this thesis were intended to obtaÍn further information on the nature and jntensity of the maintenance processes in some crop legumes (field bean, chickpea, lucerne, Pêê and kidney bean). Experiments were conducted using whole plants (grown in a naturally light glasshouse at 20oC) and mature leaves (p'lants were grov,rn for 20'40 days in a controlled environment room at ZOoC). Whole plants were used to compare main- enance coefficjents calculated using three different methods, and also to investigate the relatjonship between the respiration rate during starvation and plant dry weight. The dark respiration of mature field bean leaves was studied to determine: (i) the pattern of respiration of attached and detached leaves during a proionged dark treatment, sjnce this treatment has been used to estimate maintenance respiration, (ii) the source of carbohydrate for respiration during the dark treatment, and (iii) whether leaf senescence occurs during the treatment. The three methods of measuring the maintenance component were: (i) to allow the C0, efflux to decay in prolonged darkness to an asymptotic value which is then taken to be the maintenance value; (ii) to plot the dark C0, efflux as a function of net C0, uptake over a range of irradiances and take maintenance as the dark efflux when the net C0, uptake is zero; (iii) to pìot total C0, uPtake as a function of the growth rate and take maintenance as the COt efflux when the growth rate is zero. The maintenance coefficients so calculated for field bean, chick- pea and lucerne using methods (i) and (ii) were similar; however coefficients calculated using method (iii) r^rere significant'ly higher (p.0.05). Method ('i) was considered to provide the best estimate for maintenance because there was less opportunity for growth processes to contribute to C0, efflux. Method (ii) resulted in estimates for maintenance respiration which contained C0, efflux due to phìoem loading of assimilate'in the leaves and therefore there was greater opportunity for structuraì growth. These latter processes (phloem ioading and structura'l growth) were considered to contribute to a greater extent to C0, efflux when the maintenance coefficient was determined using method (iii). It is concluded that method (i) gives minirnal estimates of the maintenance requirements, but has the advarrtage of being simple and amenable for the mass screening of plants if those vlith a 'low'maintenance respiration are to be i denti f i eci . Maintenance respiration consurned the equivalent of 2% of the dry weight per day in field bean, chickpea and lucerne. The growth efficjency (Yn) was the same for all species (0.09 t 0.01) and was unaffected by the method of calculation. There t,úas a linear relationship between the respiration rate of whole pìants after 48 hours in the dark, and: (i) the dry weight, (ii) the total nitrogen content, and (iii) the organic nitrogen content. These relationships are inter- preted to mean that protein content vras being maintained during starvat'ion. The respiration rate of whole field bean pìants was essentia'lly constant during a 12 hour night. Defoliation and removaì of leaves from the assimilation chamber at the beg'inning of the n'ight removed 20-30% of the dry weight yet hardly affected the respiration rate of the remaining plant parts immediate'ly foìlowing defoliation. The respiration rate subsequently declined linearìy with time. This result is interpreted to mean that phìoem loading of sucrose in the leaves maintains a constant flow of assimilate to the 'sinks'of the rest of the plant. The rate of dark respiration of mature field bean leaves was dependent upon the irradiance levei during the prev'ious photoperiod. Studies on the pattern of change in dark respiration 'in prolonged darkness revealed the following: (i) Dark respiration was essentia'l1y constant during the normal 12 hour night and there was a rapid loss in leaf dry weight during this period. (ii) After 12 hours, the respiration rate decayed to anasymptotic value at about 24 hours. (i i i ) The gas exchange quotient (ul C02/vl 0r) remained above 0.90 for 60 hours when leaves were detached at the end cf the photoperiod. llhen attached leaves were measured, the quotient was above 0.90 during a normal (12h), but subsequently decljned in a comp'lex way, which indicated that protein was being oxidised. The energy cost for phloem loading and sucrose synthesìs were calculated from the Iiterature to be equiva'lent to 0.05279C respired/gC translocated. The experi- mental value, based on the difference in the respiration rate of mature leaves ciuring the normal night and after 60 hours darkness was 0.0560gC/gC.
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