Studies on the Calcicole-Calcifuge Habit: II. The
Transcription
Studies on the Calcicole-Calcifuge Habit: II. The
691 STUDIES ON THE CALCICOLE-CALCIFUGE HABIT II. THE INFLUENCE OF CALCIUM ON THE GROWTH AND ESTABLISHMENT OF FOUR SPECIES IN SOIL AND SAND CULTURES BY R. L. JEFFERIES AND A. J. WILLIS ofBristol ofBotany,University Department I. INTRODUCTION & Willis 1964) indicatesthatthe Evidencewhichhas been presentedearlier(Jefferies calcifugespeciesJuncussquarrosusL. and NardusstrictaL. are plants possessingan narrowdistribution rangein respectof levelsof exchangeablesoil nutrients, extremely especiallyregardingcalcium. Sieglingiadecumbens(L.) Bernh.is tolerantof a much widerrange,whereasOriganumvulgareL. is-restrictedto soils rich in exchangeable to assessto whatextentthe calcium.An attempthas beenmade,usingculturetechniques, in distribution patternsof theseplants are determinedby the availabilityof nutrients the soil. typesof soil in orderto study Plantsof thefourspecieshave been grownin different theirabilityto become establishedin soils fromwhichtheyare absentundernatural conditions.The plantsalso have been grownin sand culturesto whichmineralsolutions to havebeenadded suchthatthenutritional statusoftheculturesprobablyapproximated considerably conditionsfoundin naturalhabitats.As theavailabilityof calciumdiffered of in thesoilswhichhave beenanalysed,particularattentionhas beengivento theeffect in relationto the concentrationof the othernutrientson the calcium concentration establishment and growthof thesespecies. II. CULTURE TECHNIQUE Plantsweregrownunderglasshouseconditionsin polythene pots,8 in. (20 cm) deep and 6 in. (15 cm) in diameter,or else in earthenware pots of similardimensions.In orderto excludeany possibilityof nutrients beingderivedfromsourcesotherthan the culture solutions,the precautionstaken were as recommendedby Hewitt(1952). The quartz sand whichwas requiredformanyof the experiments was purifiedin a Kestnersand digester(Hewitt1952). At all timesde-ionizedwaterof specificresistancegreaterthan fourmegohms(Willis & Yemm 1961) was used in thesestudies.When the seeds were solutionwas allowedto percolatethrough sownin sand, 1 1. of the appropriatenutrient the culturein orderthatresidualacidityin-the sand shouldbe diminishedand thatthe seeds would germinate underthe same conditionsas theplantswould laterexperience. In someculturestheseconditionsparalleledthoseofnaturalhabitats.In nearlyall ofthe conditionson of nutrient experiments, plantsweregrownfromseed,so thatany effects thesestudies been used in if had couldbe elucidated; onlytransplants plantestablishment seeds of all of the Germination of thiskindwould not have been obtained. information did not here the in the fourspecieswas foundto be satisfactory temperature glasshouse; fallbelow 13?C and therewas no artificial lighting. Q JE 692 of calcium habit:influence Calcicole-calcifuge Seeds were collectedfromplants fromthe same sites each autumn.The seeds of Juncussquarrosusand of Nardusstrictawereobtainedfromplantsgrowingon theacidic peaty soil of Blackdown,Mendip (National Grid referenceST474572), of Sieglingia froma populationon the fenpeat of the Gordano valley,northSomerset decumbens limestonesoil of (ST437730) and of Origanumvulgarefromplantson theCarboniferous CadburyCamp, northSomerset(ST455722). The pots were distributedat randomin the glasshouseand periodicallymoved to theplantswereharvestedand washedin deplaces.At theend ofan experiment different weremade beforethe ionizedwaterfora shorttime;appropriategrowthmeasurements by meansof leafmaterialwas bulkedand thenanalysedforsome of themajornutrients & Willis1964). methodspreviouslydescribed(Jefferies III. GROWTH AND ESTABLISHMENT OF THE PLANTS IN A RANGE OF SOIL TYPES status To gainsomeindicationofthetoleranceoftheplantsto soiltypeand itsnutritional theyweregrownfromseed in a numberof contrastedsoils whichdo not supportthese speciesundernaturalconditions.Top soil was collectedat severallocalities,sievedand mixedbeforeitwas used (Table 1). De-ionizedwaterwas added,as required,to duplicate Table 1. The localitiesfromwhichsoils wereobtainedfor culturestudiesof thefour of thesesoils characteristics species,and some important as mgper 100g ofdry(80?C) soil. The levelsofCa, Mg,K and Na are expressed National Na Ca K Mg Soil type Gridreference Site 240 80 27 5 898 Fen peat ST437730 GordanoValley, I N. Somerset, 40 31-0 22-5 1164 Fen peat ST435729 GordanoValley, II N. Somerset, 8-5 Carboniferous 664 114-0 19-0 ST455722 CadburyCamp, limestone N. Somerset 32 5 1-5 24-0 1252 Sand-dune SS455345 BrauntonBurrows, soil N. Devon pH 36 5-2 7-4 7-9 soil cultures.Numerousseeds of Origanumvulgareand ofJuncussquarrosusweresown but in the case of Nardusstrictaand Sieglingiadecumbensonlyeightseeds weresown perpot.The plantswereharvestedin November1961aftera growingperiodof20 weeks. indicatedthatthe plantseitherdied shortlyafter The resultsof thispilot experiment or else showedpoor growthin most of the soils (Table 2). The data were germination as onlyfewplantssurvivedin each culture.Althoughthenumber nottreatedstatistically theresultssuggestedthatedaphicfeatureswerelikelyto be of soilschosenwas restricted, of considerablesignificance in limitingthesespeciesto certainsoils. The poor developmentof all of the plants,includingOriganumvulgare,in the sandy soil of Braunton ofthelow levelsofavailablenitrogenand phosphorusin Burrowsis probablya reflection in a soil thiscalcareoussand (Willis& Yemm 1961).The abilityof 0. vulgareto flourish the on has some signifibearing of pH 3-6butrichin exchangeablecalcium(fenpeat,I) and othersoil featureswhichvarywithpH, in cance of hydrogenion concentration, In thelocalitywherethissoil was collectedthewatertable influencing plantdistribution. 1959); itis possiblethatthisfeature is nearto thesurfaceoftheground(Willis& Jefferies R. L. JEFFERIES AND A. J. WILLIS 693 mayexclude0. vulgarefromsuchsoils.The growthofindividualplantsofNardusstricta variable.One of the onlytwo plants in the Carboniferouslimestonesoil was extremely the courseof theexperiment, whichsurvivedremainedat theseedlingstagethroughout whereasthe otherstayedat this stage for monthsbut subsequentlybegan to thrive. after Table 2. The averagedryweightof theshootsystemsof theplannts typesof soil for 20 weeksin different growth perplant. The figures givemgpershootsystem Juncus Sieglingia Origanum Nardus Soil type squarrosus stricta decumbens vulgare 402 Fen peat,I 49 98-0 164 Died* Fen peat,II 38 0 1-2 11-8 Carboniferous 147-0 limestone soil Died 50.0 26-0 13-0 2-0 Sand-dunesoil Died 3-1 * Fungal attack. Under naturalconditionsthe majorityof theseplantsundoubtedlywould die withina shorttimein suchsurroundings, and evenifa fewmanagedto survivefora longerperiod lead to theirdeath. fromotherspecieswould ultimately probablycompetition IV. EFFECT OF CALCIUM LEVEL ON THE GROWTH AND ESTABLISHMENT OF THE FOUR SPECIES IN SAND CULTURE 1 (1) Experiment The aim of thisexperiment, and of subsequentones describedin thissection,was to on thegermination and growthof calciumconcentrations elucidatetheeffects ofdifferent of to providesimilarconcentrations theplants.It seemeddesirablein a firstexperiment 1 Table 3. The composition of theculturesolutionsusedin Experiment 1 3 Culturenumber 6 4 5 7 mgCall. as: 4 4 4 4 4 4 4 Ca(NO3)2 0 28 92 240 442 668 4 CaCl2 32 672 8 96 244 446 Totalcalcium(mg/l.) 4 Othernutrients wereadded to all of the culturesolutions1-7 as as mg/l. follows;valuesare expressed nitrate Cu, 0 6, as coppersulphate K, 16,as potassium sulphate Zn, 0 65, as zincsulphate Mg, 18,as magnesium P, 12-2,as sodiumdihydrogen B, 3 0, as boricacid and Na, 25, as sodiumhydroxide phosphate sodiumdihydrogen phosphate Fe, 11-2,as ironcitrate sulphate Mn, 5 5, as manganese 2 the in the culturesolutionsto those foundin naturalhabitats.Accordingly, nutrients of to analyticaldata of the levels exfinalconcentrations were adopted by reference & Willis 1964) and were in soils supportingtheseplants(Jefferies changeablenutrients also similarto thevaluesgivenbyGorham& Pearsall(1956) forthemineralcomposition ofbog and fenwaters. soluAlthoughthe amountsof calcium(as CaCl2) whichwereadded to the different unaltered remained (Table 3). tions were varied,the quantitiesof the othernutrients of calcium habit:influence Calcicole-calcifuge 694 The pH of the seven culturesolutionswas adjustedto 5 0 (by NaOH); the driftenfromthe sand cultureswas of the effluent counteredin the hydrogenion concentration solufoundon all occasionsto be betweenpH 4-5and 5-5.Additionsof 100ml ofnutrient tionsweremadeweeklyto theduplicateculturesand thesandwas moistenedas required. (a) 1004 100 -0 A / E (b) 5 \ A? 75/'- 25 -u ~25- \A 0 1~10 0 500 50 100 mg Ca/I 4 10 500o 50 100 mg Ca/l levelsofcalciumsuppliedas different ofplantsin culturescontaining FIG. 1. The growth chloride.Details of thesevenculturesolutionsused are givenin Table 3; calciumconis shownon a log scale. Mean dryweightsof theshoots(a) and roots(b) of centration *; vulgare, 0; and Origanum decumbens, *; Sieglingia *; Nardusstricta, Juncus squarrosus, are plotted. Table 4. Analysesof leaf tissueof Juncussquarrosusand dryweightsof theplants in cutlture experiments 3 as in all theseculturesall theplantsdied. No resultsare includedforExperiment mgCa/l. Plantsofnaturalhabitats Minimum Maximum Mean 1 Experiment Culture1 Culture2 4 8 2 Experiment Culture1 Culture2 Culture3 Culture4 4 8 32 96 4 Experiment Culture1 Culture2 12 24 drywt. mg/g Na Mg K 0-3 0-9 063 1.1 2-3 1-5 100 18 6 150 1-8 6-3 10-7 4-6 Sampleslost Ca 1-7 59 2-5 6-5 6-3 6-9 24-0 -65 130 122 12-6 10-8 Meanwt./shoot Mean wt./root of (mg) of (mg) (O% 45 41 100 91 32 27 100 84 51 57 41 22-5 89-5 100 72 39 5 415 58 13 7 71 100 22 12 10 7-3 100 73 6 5 100 83 max.) (O% max.) 05 1-9 10 2-6 3-9 7-8 5-5 Not enoughmaterial foranalyses vulgareweresownbutthenumber and of Origanum NumerousseedsofJuncussquarrosus was limitedto sixteenand eight of seeds of Nardusstrictaand of Sieglingiadecumbens was begunin May 1960and theplantswereharvestedaftera The experiment respectively. periodof 28 weeks. R. L. JEFFERIES AND A. J. 695 WILLIS The resultsof the growthof thesefourspeciesexpressedin termsof dryweightare in survivaland in growthmade shownin Fig. 1, and also in Tables 4-7. The differences by the plantsin the cultureshavingvariouslevelsof calciumare evidentfromFig. 1. affect thegrowthofthe Althoughtheresultssuggestthatthedifferent culturetreatments Table 5. Analysesof leaf tissueof Nardus strictaand dryweightsof theplantsin culture experiments withinan experiment Wherethedryweightsoftheplantsgrownunderthevariousculturetreatments different difference is shown. are significantly (5 % level)theleastsignificant (L.S.D.) fortheexperiment mgCa/l. Plantsofnatural habitats Minimum Maximum Mean Ca mg/gdrywt. Mg K Na Mean wt./shoot (mg) (% of max.) 09 16 13 09 1-8 14 50 11-5 79 0.1 08 03 1-6 69 1-6 93 1-3 103 83 07 4-1 39 1 Experiment Culture1 Culture2 Culture3 Culture4 Culture5 4 8 32 96 244 0-6 1D1 2-4 4-8 11-7 29 2-8 3-2 2-8 26 9-2 11-8 12 8 15-0 10 4 2 Experiment Culture1 Culture2 Culture3 Culture4 Culture5 L.S.D. 4 8 32 96 244 0-8 10 2-0 30 27-9 30 3-1 3-2 19 4-8 76 70 52 4-3 6-8 3 Experiment Culture3 192 4-4 3-4 12 2 1-7 112 4 Experiment Culture1 Culture2 Culture3 L.S.D. 12 24 96 2-3 22 4-7 37 3-1 2-5 8-3 100 70 5 Experiment Culture1 Culture2 Culture3 Culture4 Culture5 Culture6 Culture7 Culture8 L.S.D. 04 08 32 9-6 24-4 44-6 67-2 89-6 3-7 3-5 27 2-9 7-0 9-6 12-5 12-4 41 1-2 1-3 1-7 1-7 1-5 2-8 2-3 5-4 10-3 5-8 3-4 1-4 2-4 3-2 1-2 67 90 100 80-5 38 Mean wt./root (mg) 35 51 58 27 9 No. of (% of seeds No. of plants max.) sown harvested 60 88 100 47 15 16 16 16 16 16 12 15 16 9 1 26 32 70 100 15 15 15 15 15 15 12 9 12 10 3 100 12 3 2-0 37 1-4 88 1-7 163 80 22-5 54 100 12 12 12 11 12 7 2-1 50 8-9 25 50 49 0-8 97 0-8 93 0 4 210 1-6 135 0-8 235 107 21 105 21 41 39 89 57-5 100 10 10 10 10 10 10 10 10 5 8 5 6 9 6 4 4 09 07 04 11 16 6 56 77 195 301 74 138 18 5 58 25 5 70 65 155 220 100 24-5 32 88-5 plantsto a considerableextent,statisticaltestsmade by means of thevariance-ratio (Ftest)and multiplerangetest(Duncan 1955)showedthat,in respectofmeandryweightsof and Nardusstricta,differences shootsand rootsof Sieglingiadecumbens from resulting in theplantswhichsurvived,did not quitereachsignifithevariousculturetreatments, cance at the 5% levelof probability. However,in thisexperiment onlya relatively small of calcium habit:influence Calcicole-calcifuge 696 Table 6. Analysesof leaftissueof Sieglingiadecumbensand dryweightsof theplantsin cultureexperiments withinan experiment oftheplantsgrownunderthevariousculturetreatments Wherethedryweights is shown. (L.S.D.) fortheexperiment difference (5 % level),theleastsignificant different are significantly mgCa/I. Plantsofnatural habitats Minimum Maximum Mean Ca mg/gdrywt. Na K Mg 09 40 19 04 2-8 1-3 50 14 4 101 03 55 19 1-8 19 1-6 18 19 90 1 Experiment Culture1 Culture2 Culture3 Culture4 Culture5 Culture6 4 8 32 96 244 446 1P7 2-0 47 95 195 14 3 59 5-5 50 40 46 3-1 15 3 119 124 12 1 129 15 1 2 Experiment Culture1 Culture2 Culture3 Culture4 Culture5 L.S.D 4 8 32 96 244 1-2 21 40 70 23 5 61 49 50 3-5 39 76 75 93 7-3 56 3 Experiment Culture1 Culture2 Culture3 24 48 192 69 5-8 119 53 38 4-5 4 Experiment Culture1 Culture2 Culture3 Culture4 L.S.D. 12 24 96 288 5 Experiment Culture1 Culture2 Culture3 Culture4 Culture5 Culture6 Culture7 Culture8 Culture9 L.S.D. 04 08 3-2 96 24 4 44 6 67-2 89-6 131-2 No. Mean Mean of No. of wt./root wt./shoot (mg) (% of (mg) (% of seeds plants max.) sown harvested max.) 70 100 96 5 945 93 66 5 80 127 96 100 31 39 63 100 76 79 24 31 8 8 8 8 8 8 7 6 8 6 4 7 1-5 19 04 71 2-3 112 2 9 310 6-4 46 40 6 22 36 100 14 22 101 138 253 30 37 9 40 55 100 12 12 12 12 12 12 10 11 11 9 7 16 9 13 2 17 2 2 1 24 1-8 101 19 34 23-5 100 33 5 8 8 8 1 5 1 54 19 3-7 3 9 11 9 12-8 58 550 13 3 105 63 3.3 1 9 157 08 196 11-4 151 8 3 72 92 80 100 77 365 8 8 8 8 6 6 5 4 11 2 1 1-8 4 5 2-5 07 4-8 1 8 9*9 2 2 12 0 2-3 11 142 16 7 4 4 22-7 10 1 53 30 4.7 4-5 3-4 2-6 30 5-1 10 1 6 6 6 6 6 6 6 6 6 3 4 3 4 4 2 2 3 1 1-3 10 07 07 08 03 06 19 19 99 141 136 133 131 94 41 89 52 24 12-5 27 58 125 144 66 5 86 186 216 100 61-5 133 34 73 56 5 R. L. JEFFERIESAND A. J. WILLIS 697 numberofplantswas studied.In additionto valuesof dryweights,resultsof analysesof as wellas thedata of some subsequentones,are summarleaf tissuesin thisexperiment, ized forconveniencetogetherin Tables 4-7. Here also are includedthevaluesforleast differences, in dryweightsof shoots and roots of plants grownunderthe significant different levelsofcalciumsupply,calculatedbythemultiplerangetestprocedure(Duncan were in whichdifferences on N. strictaand Sieglingiadecumbens 1955) forexperiments oftheplantsinculture Table 7. AnalysesofleaftissueofOriganumvulgareanddryweights experiments mgCa/I. Plantsofnatural habitats Minimum Maximum Mean mg/gdrywt. Ca Mg K Na 9-8 23.1 16-5 14 80 4-3 9-2 28.8 16-5 0.1 04 03 Mean wt./shoot Mean wt./root (mg) (% of (mg) (% of max.) max.) 1 Experiment Culture3 Culture4 Culture5 Culture6 Culture7 32 96 244 446 672 9.1 11 1 13-0 26 2 29 5 85 25 3 65 56 17 5 16 9 5-4 Sampleslost 05 0o4 08 07 11 22 57 75 72 14-5 29 76 100 96 14 20 40 53 48 26 38 75-5 100 90 5 2 Experiment Culture3 Culture4 Culture5 Culture6 Culture7 Culture8 Culture9 32 96 244 446 672 896 1312 2-8 57 316 19.1 53 37-1 69 6 40 4-2 52 5.9 40 2-6 4-1 8.4 44 4-3 43 4'3 4-7 94 02 0.1 17 1.1 1.5 07 2-0 45 39 88 164 143 130 37 27 5 24 53-5 100 87 79 22-5 24 18 36 52 42 32 12 46 35 69 100 81 62 23 3 Experiment Culture3 192 4 Experiment Culture2 Culture3 Culture4 24 96 288 5-4 14 2 13-7 4-4 45 3'0 7-8 24 14 5 Experiment Culture5 Culture6 Culture7 Culture8 Culture9 24 4 44-6 67-2 89-6 131-2 25 7 29 2 38-0 62 7 45-7 26-7 19-7 21-5 240 15-0 5.3 48 8-5 50 3-8 11 100 5 100 1.0 1-5 16 115 102 120 96 85 100 39 54 29 72 100 54 9-6 45 9.1 100 12-5 30 40 40 56 76 39 5 52 5 52-5 73-5 100 27 32 37 42 58 46.5 55 64 72-5 100 Not enoughmaterialfor analyses at leastat the 5% level.No statisticalanalyseswerepossibleof theresultson significant in separating Juncussquarrosusand Origanumvulgare,partlybecause of the difficulties same cultures. the in the completely rootsof theplantsfromthoseof others was In Experiment1 in culturespossessinghighcalciumconcentrations germination cultures in such or did not to else occur. Seeds of failed Juncus germinate squarrosus delayed are ofseedsof Origanumvulgarewas retarded.These observations whilethegermination in generalagreement withthefindings of Webb & Hart (1945). Whereas0. vulgareand, at thesehighcalcium to a lesserextent,Sieglingiadecumbens werecapable of surviving 698 of calcium habit:influence Calcicole-calcifuge and becomingestablished,Juncussquarrosusand Nardusstrictaquickly concentrations was fixedabout 4 weeks after succumbed;consequentlythe course of the experiment conductedwhichinvolvedthe This was evidentin all of the experiments germination. growthof plantsfromseeds. Juncussquarrosusand, to a lesserextent,Nardusstrictawerelimitedto the cultures resultsthat whichcontainedlow calciumlevels; it would appear fromtheexperimental Juncussquarrosusis a strictercalcifugethan Nardus stricta.This is consistentwith madein naturalhabitats,foralthoughNardusis capable ofgrowingat very observations highercalcium low calciumlevelsitwas foundmoreoftenin soilswhichhavesubstantially & Willis1964). levelsthanthosesoilsin whichJuncussquarrosusis established(Jefferies Origanumvulgarecannotsurvivein theculturespossessinglow calciumconcentrations. In such culturesthe seedlingswere small and stuntedand showedcalciumdeficiency symptomsin theirleaves and at the growingpoints.Sieglingiadecumbenssurvivedat in highercalciumlevelsthan Nardusstricta,whichis in keepingwithits distribution naturalhabitats. is that with increasingcalcium concentration A notablefeatureof this experiment therewas usuallypoorerroot development.This observationof deleteriouseffectsof of Olsen (1942), withthefindings highcalciumlevelson growthis in generalagreement Webb & Hart (1945) and Rorison(1960). N. strictaat thehighercalciumlevelsformed oflateralswhereasat thelowercalciumlevels shortstuntedrootswithpoor development therewas a profusedevelopmentof root lateralsforminga mattedgrowth. ofthesespeciesin thevariousculturesare in growthand establishment The differences withtheresponseoftheplantsto thewidevarietyof soil conditionsin natural consistent the quantityof nutrientsin the shoot tissuesof each of these habitats.Furthermore, speciesshowedsome parallelismto thatin the plant materialcollectedfromthe field is clear. (Tables 4-7). In the case of Juncussquarrosusand Nardusstrictathe similarity However,in cultureswherethe levelof calciumis highand the growthof theplantsis fromplants of naturalhabitatsin respectof poor, thereis a considerabledifference in thetissues,especiallyof calcium.The same observationis applicamountsof nutrient able to Sieglingiadecumbens.At low calcium levels an accumulationof monovalent takesplace. cationsin thetissuesof all fourspecies,but especiallyofJuncussquarrosus, Magnesiumcontentis also highin theshootmaterialofall ofthespeciesbutit decreases withincreasedcalciumintheculturesolutions.Theseobservations in Sieglingiadecumbens King & Handley(1950), are in agreementwithearlierfindingsof Jacobson,Overstreet, Epstein(1961), Snaydon& Bradshaw(1961) and others.Thereis no positiveindication thatincreasingthe calciumcontentof the fromthe analyticaldata of thisexperiment culturesolutionresultsin a decreasein the uptakeof potassium,exceptpossiblyin the case of Origanumvulgare(Fig. 7). However,becauseofpoor growthofplantsin cultures betweenions duringsaltuptake,considerable whereone mightexpectseverecompetition of interpretation. analyticalproblemsariseas well as difficulties 2 (2) Experiment was similarto Experiment1, exceptthatcalciumnitratewas the sole This experiment sourceofcalcium,calciumchloridebeingreplacedbythenitrate.Bychangingtheanionit was possibleto exploreto whatextentthechlorideion accountedfortheresultsobtained The procedureadoptedwas as previouslydescribedexcept in theprecedingexperiment. thattherangeof thecultureswas increasedto includetwo additionalhigherconcentrations of calcium.Cultures1 to 7 were of the same calciumlevelsas in Experiment1, 699 R. L. JEFFERIES ANDA. J. WILLIS 896 and 1312mgCa/l.The amountsof other butcultures8 and 9 contained,respectively, nutrientsin the culturesolutionswere as shownin Table 3. The seeds were sown in March 1960 and theplantsharvestedin August1960. The results,includedin Tables 4-7, are essentiallysimilarto those givenearlierin squarroindicating theinabilityoftheplantsto growin particularcultures(Fig. 2). Juncus highcalciumlevels, sus and Nardusstrictafailedto becomeestablishedin sandcontaining as did Sieglingiadecumbens.In cultureswhich containedsmall amountsof calcium, Origanumvulgarewas unableto surviveand theleaves showedseverecalciumdeficiency as well as nitrogendeficiency. at thelow calciumconcentraThe growthof Nardusstrictaand Sieglingiadecumbens low (Tables 5 and 6). This poor growthwas also tions was foundto be significantly forin thepreviousexperiment probablya reflection of thelow levelof availablenitrogen, (a) 100 E 75 AU (b) 100U '. 75 / O / 450 / L75 0 4 10 50 . . ~~~~~~~~~25~~ in 50 100 mg Ca/L / 500 1000 0 4 10 50 100 mg Ca/l 500 1000 levelsofcalciumsuppliedas different FIG.2. The growthofplantsin cultures containing was nitrate.The graphis constructed in the same way as Fig. 1, but the experiment extended to twoadditionalhighlevelsofcalcium(cultures8 and 9). Juncus squarrosus, @; vulgare, v. Nardusstricta, A; Sieglingia decumbens, 0; Origanum wherethenitrogen levelswereconstantthegrowthmadebythesetwo specieswas similar The analysesofplant and substantialin culturescontaininglow calciumconcentrations. tissue(Tables 4-7) also yieldedsomewhatsimilarresultsto thoseof thepreviousexperimentbut again indirectevidenceof ion antagonismduringsaltuptakeis lacking.In all fourspecies,a highconcentration of calciumin theculturesresultedin largeamountsof calciumin theleaftissue. havebeenfoundin naturalhabitatswhere, and Origanum vulgare AlthoughS. decumbens such in the soil, the ratio of exchangeablecalciumto the otherexchangeablenutrients as potassiumand sodiumis high,at similarratiosin theculturestheplantsdied or else of calciumwas different from showedpoor growth.It is probablethattheconcentration thatexperiencedunderfieldconditions,as discussedlater,and thisfeaturemayaccount forthe resultsobtained.However,thepossibilityexiststhatsuch a speciesas Sieglingia decumbens may be composedof a numberof distinctpopulations,some of whichare establishedon chalkand limestonesoils.Whenseed of capable of becomingsuccessfully Sieglingiawas used whichhad been collectedfromBrauntonBurrows,a highlycalcareoushabitat,thegrowthof theseplantsin sand culturesgave similarresultsto those alreadypresented.Some explanationotherthan the existenceof distinctpopulations of calcium habit:influence Calcicole-calcifuge 700 soughtto accountforthe inabilityof thisgrassto growin culturesconwas therefore indicatethatedaphic taininghighlevelsofcalcium.The data presenteddo notnecessarily workis required do not occurand clearlymoreexperimental ecotypesof S. decumbens involvingseedscollectedfroma largenumberofhabitatsin orderto solvethisproblem. 3, 4 and 5 (3) Experiments in whichtheculturesolutionapplied on experiments The resultsgivenabove are based exceptforthe level of the of nutrients to the sand containedthe same concentration to the othercations in these calcium ratio of the particularcalcium salt. Although (based on investigated were solutionswas similarto the ratios found in soils which E100 EXPT.4 EJ I ~~~ 50 12 Plants died 50 I. 500 100 mg Ca/l 1000 3000 1 I EXPT. 2 ~~~~~~~~~~~~~~~~~~Plants . I 4 10 idied 100 50 mg Ca/L 500 1000 levelsof calcium. different in culturescontaining squarrosus FIG. 3. Growthof Juncus The meandryweightoftheshootsis shown. exchangeableion levels),some of the speciesfailedto growin certaincultures,as disof mineralsin cussedearlier.This mayhave been caused,in part,by the concentration the sand culturesnot beingthe same as that experiencedby the roots undernatural conditions.The resultsofthesoil analysesare based on unitweightand suchdata maybe misleadingifa numberof soil typesare beingcompared(Fried & Shapiro1961; Jefferies & Willis 1964). Plants died Plnts EXP died ro E 100 24 500 1000 5000 4 10 50 100 500 1000 0 0 FG0 EXPT.4 . 1eXPTa 50- cm mg Ca/L mg Ca/L different levelsof calcium. FIG. 4. Growthof Nardusstrictain culturescontaining were set up whereadditionsof nutrients To investigatethis problem,experiments weremade based on thesameratiosofcalciumto otherions as previouslyemployedbut was changedfromthatoftheearlierexperiments ofall ofthenutrients theconcentration werechosenwhichwere,in respectof all of by the same factor.Three concentrations 701 ANDA. J. WILLIS R. L. JEFFERIES the nutrients,six times (Experiment3), three times (Experiment4), and one-tenth involvingcalciumnitrate used in the experiment (Experiment5), of the concentrations (Experiment2). However,as in Experiment1, calciumchloridewas the main calcium sourceand not calciumnitrate.In each of the Experiments 3-5, nine culturesolutions 1002 EXPT1 EXPT.1 3 00151 T< ii 50-- ro 0 0 ~~~~~~~~~~EXPT.4 100 o 24 mq Ca/l 30000 4 1000 500 5000 0 10 E XPT. 5 50 500 1000 0 mg Ca/ 1I 04 3000 10 10 0 decumbens in culturescontaining different FIG. 5. Growthof Sieglingia levelsof calcium. I were10 rangedfrom7872 mg/I.in culture9 wereemployed;thetotalcalciumcontenttherefore procedure of Experiment3 to 0 4 mg/l.in cultureI of Experiment5. The experimental was as previouslydescribedbutbecause ofthelargenumberofpotsrequiredforreplicathe at thesame time;consequently tionit was not possibleto startall threeexperiments resultsare not directlycomparableand can be expressedon a relativebasis only. 100[ EXPT. 3 It I9 died Plants E @ 24 ~100 Plants did 12 50 500 ! 100 4 10 50 4 1 5 10 mg Ca/l Ilill 500 1000 0 100 sdied |PLant 100 5000 1000 I EXPT. 2 500 mg Ca/l 50 00 levelsof calcium. of Origanumvulgarein culturescontainingdifferent FIG.6. Growth in Juneand in May 1960and all 3 and 4, seedsweresown,respectively, In Experiments 5, seedsof Nardusstricta of theplantswereharvestedin December1960.In Experiment weresown in September1960,and the plantsharvestedin decumbens and of Sieglingial vulgarewere sown in September1960 and the plants July1961; seeds of Origan7um of calcium habit:influence Calcicole-calcifuge 702 harvestedin December1960; and seeds of Juncussquarrosusweresownin August1961 was undertaken. but no detailedharvesting The growthmade bytheplantsis shownin Figs. 3-6. Statisticaldata, dryweightsand in the leaf tissuesare givenin the resultsof the analysesof variousmineralnutrients Tables 4-7. The resultsindicatethatthepatternof behaviourof all fourspeciesin these conditionsis moreproculturesis essentiallysimilar,althoughtheresponseto different of all of the nutrients nouncedin some of the species.An increasein the concentration of the plantsto cultureswherethe ratio of calciumto the other leads to the restriction in althoughjust surviving cationsis low,as foundin a non-calcareoussoil.J. squarrosus, diesin all cultures ofnutrients, suchcultureshavingthreetimestheoriginalconcentration of Experiment1. The establishment of nutrients whichhave six timesthe concentration restricted in culturesis likewiseprogressively of Nardusstrictaand Sieglingiadecumbens Origanumvulgarebehavesin a similarmanner, ofnutrients. thehighertheconcentrations wheretheratioofcalciumcomparedto onlyin culturesofhighconcentrations surviving is low. othernutrients 60 /a a) ' 40A * 15 30 10(b) 20 5; C:20 100 50 mg Ca/l of culture solution 10 FIG. 2 50 \ E 10=== 30 50 mg Ca/l 300 100 of culture solution 450 7. Thechangesinlevelsofcalcium(0), magnesium (A) intheleaves (o) andpotassium calcium of (a) Sieglingiadecumbensand (b) Origanumvulgrare grownin culturesof different levels. For Sieglingia decumbensthe levels of magnesiumand potassium were constantin the culture solutions at 54 and 48 mg/I.respectively(Exp. 4). For Origanumvulg'arethe levelsofmagnesiumand potassiumwereconstantin theculturesolutionsat 18 and 16 mg/I. respectively(Exp. 1). Inabilityto survivein such culturesdoes not appear to be an osmoticeffectas the culturesolutionis onlyaboutfiveatmospheres. osmoticpressurein themostconcentrated is protracted.Poor growthof but further The seeds germinatesuccessfully development amountsof nutrientsin the plantsin theseculturesis associatedwithwidelydiffering leavesas comparedwiththoseofplantsin naturalhabitats.An increasein calciumin the mineralsolutionsnot onlyleads to a greateraccumulationof calciumin theplanttissue and potassium(Tables4-7). butitis also oftenassociatedwitha lowerlevelofmagnesium This is especiallyevidentin Sieglingialdecumbensand in Origalnum vulgare(Fig. 7). Whethertheseresultsrepresentactiveaccumulationor just Donnan exchangereactions is not known. 2 thereis a relatively of Experiment oftheconcentrations In cultureshavingone-tenth widerangeof tolerancebyall fourplants;heresomeplantsbecomeestablishednot only is low but also in cultures in cultureswherethe ratio of calciumto the othernutrients whereit is similarto thatfoundin a calcareoussoil. Nardusstrictaand Sieglingiadecumis high.The wheretheratioofcalciumto theothernutrients bensgrowmostsuccessfully R. L. JEFFERIES AND 703 A. J. WILLIS conditions underexperimental resultsindicatethatit is possibleto growS. decumbens is similarto thatin a calcareoussoil. wherethe ratio of calciumto the othernutrients Nardusstrictabehavedin a similarfashion,yetwiththe exceptionof a fewcontinental localities(Klika 1936) thereare no recordsof thisplant growingon limestonesoil. It in theseparticularsand cultures ofnutrients is probable,however,thattheconcentration involvingJuncus is low comparedwiththatof calcareoussoils.Althoughtheexperiment theconcentraitwas clearthat,evenin culturesof one-tenth was discontinued squarrosus 2, theplantgrowsonlyat thelow calciumlevels,dyingin therestof tionsof Experiment of indicatethat,dependingupon the concentration the cultures.The resultstherefore it is possibleto growat leastthreeofthesespeciesin a rangeof cultureswhere nutrients, considerably. differs theratioof calciumto the othernutrients V. THE EFFECT OF CALCIUM LEVEL ON TRANSPLANTS 6 Experiment to plantsgrownfromseed ratherthanto refer described previously of results All the to discoverhow far similar undertaken was A experiment pilot of transplants. studies resultsto thosegivenabove could be obtainedwhenvigorousyoungplantsweretransculturesolutions. ferredto different of Nardus strictaand Sieglingia Table 8. Dry weightsof transplants calciumlevels decumbensgrownin culturesof different and the average Four plantsweregrownundereach treatment (g) ofshootsand rootsperplantare given. initialandfinalweights Culture a b c Total calcium (mg/l.) 32 446 1312 Initialweightof transplants Nardusstricta Shoot Root 3-2 4-8 3 75 4-2 3-0 1.15 0 40 0 37 Sieglingia decumbens Root Shoot 1-38 1.5 055 105 0-25 05 0-32 0-20 weregrownfromseed in cultures Plantsof Nardusstrictaand Sieglingiadecumbens was low (thesolutionwas thatused for wheretheratioof calciumto theothernutrients in October1960,after culture3, Experiment1) and thensome of themweretransferred a growingperiodof4 months,to cultureswheretheratioofcalciumto theothernutrients was high.In all, the effectof threecultureregimeswas investigated:culturea was the culturein whichtheplantswereraised,b was thesameas culture6 of Experiment1, and c was similarto culture9 of Experiment2 exceptthat calciumwas presentchieflyas the chloride.The levels of calciumare givenin Table 8. From October1960theplants weretreatedwiththe appropriateculturesolutionsat the rateof 100 ml per weekuntil was terminated. August 1961 whenthe experiment of both speciessurvivedin the It is evidentfromtheresults(Table 8) thattransplants showedthat cultureswhichcontainedhighcalciumlevelsalthoughearlierexperiments not were nearlyso theseedlingsdiedin suchcultures.This indicatesthattheadultplants the Moreover cultures. the in sensitiveas the seedlingsto thelevelsof mineralnutrients here and kind reported of the onlyin studies resultsshowthedangerof usingtransplants 704 of calcium habit:influence Calcicole-calcifuge solelyon thedata obtained.Thispointhas beenstressed basingecologicalinterpretations by Rorison(1960). VI. DISCUSSION The experiments indicatethatit is possibleto growthe fourspeciesstudiedundernutritionalconditionswhichapproximately parallelthosefoundin theirnaturalhabitats, and theevidencesuggeststhattheconcentration of calciumrelativeto theconcentration plantdistribution. of othernutrients in the soil may be an importantfactoraffecting Analyses of the soil supportingJuncussquarrosusshowed that the exchangeable & Willis measured(Jefferies calciumlevelis similarto the levelsof the othernutrients was probablyfar of nutrients 1964). Exceptin Experiment5, wherethe concentration belowthatfoundin naturalhabitats,J.squarrosus failedto becomeestablishedin cultures whichpossessed a fairlyhighratio of calciumto the othernutrients.Under natural are normallyfoundin mineral conditionshighratiosof calciumto the othernutrients of nutrients in the soil solutionis probablymuchhigher soils; herethe concentration thanthatin mostorganicsoils and theplantdoes not surviveat theseratios. Nardusstrictadoes not appear to be such a strictcalcifugeas Juncussquarrosus.It growssuccessfully in naturalconditionsnot onlywheretheratioof calciumto theother conis 1: 1 but also whereit is as muchas 7: 1. Underexperimental major nutrients of all ditionsthisplantwas capable ofgrowingat suchratiosprovidedtheconcentration was nottoo high. nutrients Sieglingiadecumbens,althoughmore prevalentin non-calcareoussoils, is found growingon chalkand limestonesoilswheretheratioofcalciumto each oftheothermajor nutrients is extremely high.Again,bychoosingappropriate measured,exceptmagnesium, it was possibleto growthisgrassin cultureshavingratios concentrations of nutrients, of calciumto theothernutrients in naturalhabitatswhereit similarto thoseexperienced flourishes. ofthedistriin an understanding The resultsobtainedare probablyimportant butionof thisplant.S. decumbens showsmarkedlyhighabundanceon chalksoilsin the wettestregionsofthewestern chalkin Britain(J.F. Hope-Simpson,personalcommunication). Whereit growson the chalk in East Angliaand south-eastEnglandthesitesare generallyeitherchalk heaths(Tansley 1949) or else the localitieshave a highhumidity in thewesternhalfof thiscountrypossibly (Perring1960).The highrainfallexperienced limestoneand chalksoils and leads to a leachingof the surfacelayersof Carboniferous of thusa loweringoftheconcentration ofnutrients in theselayers,whilethedevelopment chalkheathin theeasterncountiesproducesa similarresult.Undertheseconditionsthe to allow S. decumbensto concentrationof nutrientsis possiblylowered sufficiently colonizesuchsitesand becomeestablished.Howeverit wouldappearthattheconcentrationis not low enoughforNardusstrictato establishitselfin thesehabitats. ofSieglingia Similarconsiderations maybe advancedto explain,inpart,thedistribution resultsof a decumbenson Carboniferouslimestoneand sand-dunesoils. Preliminary surveyof calcareoussoils in whichthisgrassgrowsindicatethatsitesbearingtheplant have soils in whichthe humusand watercontentare highrelativeto those of soils of comparablesitesfromwhichitis absent.Thereare also indicationsthatin localitieswhere of the other the plant growsat fairlyhigh calciumconcentrations the concentrations in thewintermonths. are also high.Seeds of S. decumbens nutrients appearto germinate Duringthisperiodthereis heavyrainfalland poor evaporation,and probablyconsiderlow calciumconable leachingof the surfacelayerstakes place, leadingto a relatively centration. ANDA. J. WILLIS R. L. JEFFERIES 705 evidence oftheseedling.Experimental The discussionabove refersto theestablishment suggeststhatwhentheplanthas reachedtheadultstageitis able to toleratemoreextreme conditions.There is some evidencethat the roots of matureplants of S. decumbens calcareous rock material.Whethersuch roots are actively are capable of penetrating ofions intotheplantis unknown.Howeverin theobservations involvedin thetransport madein thepresentstudy,therootsystemofthisspecieswas foundto be confinedto the surfacelayersin calcareoushabitats. Origanumvulgareis a calcicoleplant; the resultsfromExperiments1 and 2 indicate ofcalciumdeficiency. thatitis unableto growat low calciumlevelsand showssymptoms 2 theplant of culture9 in Experiment Howeverat the veryhighcalciumconcentration poor growth.Thismayhave beenbecausetheconcentration was foundto showrelatively undernatural of calciumin thesecultureswas higherthanthatwhichtherootsexperience conditions.However0. vulgarehas beenregardedas a 'marginalcalcicole'(Praeger1950; in part of its poor responseto Clymo 1962)and the presentresultsmay be a reflection veryhighcalciumconcentrations. calcium Althoughsimilarresponsesof each of thefourplantsare shownat different levels,theresultssuggestthesame patternof behaviourof thefourplantsto alterations in thecalciumlevelsin thenutrient solutions.This poses the questionas to the nature of the mechanismsgoverningtheseresponses.It is postulatedthatcalcicoleplantsare ions whichare considerablymoreselectivethancalcifugeplantsregardingthe different low net in calcicole plantsmay involve,in part,a relatively absorbed.This selectivity uptake of calciuminto the tissuewhen theseplants are grownin calcareoussoils. In and such plantsgrowingunder may not be so efficient calcifugeplantsthis selectivity highcalciumregimesabsorbconsiderableamountsofcalciumintothetissueat theexpense of othercations. In a comparativestudyof thegrowthof fourspecies,Sinapisalba, Dianthusbarbatus, of Hordeumdistichumand Tussilagofarfara,in solutionsof different concentrations calcium,Olsen(1942) foundthat,exceptforT.farfara,increasedcalciumlevelsresultedin diminishedamountsof potassiumand magnesiumin the leaves. In contrast,in the calcicoleT. farfara,increasedcalciumin thenutrient solutionled to increasedabsorptionof in theabilityof rootsto absorbmaybe potassium.It is possiblethatsimilardifferences involvedin ironuptake,and mayexplainin partthephenomenonofironchlorosiswhich is oftenobservedwhencalcifugeplantsare grownin calcareoussoils (Olsen 1923 and others),but here iron availabilitymust also be an importantfactor.This differential may occur at the root surfaceor at certainsiteswithinthe plant. The very selectivity rapid deathof seedlingsof calcifugeplantsin culturescontaininghighlevelsof calcium as beingsufficient to accountforcompletefailureso does not suggestsimpledeficiency If thereis a largeuptakeof calciumintothe tissue,unlessit is soon aftergermination. to an inactivecentreor precipitated as oxalateit maybringabout inhibition transported of enzymeaction. Dixon & Webb (1958) quote considerableevidencebased on in vitro studiesthatcalciumcan stopthefunctioningof enzymesystemsbecauseit competeswith othercationssuchas manganeseand magnesium forco-factorsitesofparticularenzymes. Origanumvulgarewhen grownat low calcium levels dies fromcalcium deficiency. However,it seemsunlikelythatcalcicoleplantsrequiremore calciumforgrowth(per unitweightofprotein)thancalcifugeplants.Analysesofplantswithsimilargrowthhabit, such as Juncussquarrosusand J. inflexus, indicatethatthe calciumlevelin thetissuesis similarin bothplants,althoughone is a calcifugeand the othersomewhatcalcicolous. for calciumin calcicole If the mechanismof ion accumulationinvolvesa low affinity 706 of calcium habit:influence Calcicole-calcifuge is also low thenetuptakeofcalcium plants,thenin soilswherethecalciumconcentration of thiselementin the plants. may be smalland resultin deficiency The precedingdiscussionrelatingto salt uptake is clearlyhypotheticaland several could be advanced.The aspectsconsideredare, however,very interpretations different Althoughstrictcalcifugesand calcicoleshave in relationto plantdistribution. important narrowlimitsoftoleranceof calciumcontentofthesoils,manyplantshave a fairlywide in the responseof these in respectof calciumlevels.Differences rangeof distribution concalciumlevelsmaynot be readilyapparentunderexperimental speciesto different factorswhich in soilsare certainly and availabilityofnutrients ditions,butconcentration Here considerableemphasishas been given requireconsiderationin plantdistribution. oftheothernutrients in relationto theconcentration ofcalciumconcentration to theeffect and growthof the speciesstudied,but undoubtedlymanyother on the establishment habit. of thecalcicole-calcifuge nutritional factorsare also involvedin determination SUMMARY of the establishment and growthof fournativeBritish investigation An experimental and Origanumvulgare) species(Juncussquarrosus,Nardusstricta,Sieglingiadecumbens habit.The plantshavebeen to thecalcicole-calcifuge has beencarriedout withreference conditionsofthe in cultureswhichprobablyparalleledthenutritional grownsuccessfully soils of theirnaturalhabitats. relative Particularattentionhas been givento theinfluenceof calciumconcentration of the plants. on thegrowthand establishment oftheothernutrients to theconcentration were concentration responsesofthefourspeciesto calcium Well markedand distinctive of levels low and Nardusstrictagrewonlyin culturescontaining shown.Juncus squarrosus of calcium. at Sieglingia vulgarewas mostvigorous highlevels calcium,whereasOriganum on theotherhandwas able to toleratea widerangeof calciumconcentrations. decumbens werelow in theculturesolutions,it Providedthatthelevelsof all ofthemineralnutrients at was possibleto growNardusstrictasuccessfully highratiosofcalciumto othercations, werehigh,even Origanumvulgaredid not thrive but whenthelevelsof all thenutrients at thesehighratios. in the shoottissueof plantsgrownin culturesoluThe amountsof mineralnutrients valuesobtainedfromplantsofnaturalhabitats. tionswerecomparedwithcorresponding conditionsof the culturesverydifferent Whenpoor growthwas made undernutritional in the fromthose of the normalhabitatsof the plants,the levelsof mineralnutrients naturally. growing of plants fromthose different leaves weresubstantially and oftheresultsofthesestudieson thedistribution The broadecologicalimplications growthof thefourspeciesare discussed. REFERENCES approachto partofthecalcicoleproblem. J. Ecol. 50, 707-31. Clymo,R. S. (1962).An experimental Dixon,M. & Webb,E. C. (1958).Enzymes.London. F-tests. Biometrics, 11, 1-42. Duncan,D. B. (1955).Multiplerangeand multiple cationtransport byplantcells. PlantPhysiol. roleofcalciuminselective E. (1961).Theessential Epstein, 36, 437-44. inionuptake. Annu.Rev.PlantPhysicl. 12, Fried,M. & Shapiro,R. E. (1961).Soil-plantrelationships 91-112. ofsomebogand andcalciumcontent conductivity specific E. & Pearsall,W. H. (1956).Acidity, Gorham, Britain. J. Ecol. 44, 129-41. fenwatersin northern R. L. JEFFERIES AND A. J. WILLIS 707 methods usedinthestudyofplantnutrition.Tech.Commun. Hewitt, E. J.(1952).Sandandwaterculture Bur.Hort.,E. Malling,22. R., King,H. M. & Handley,R. (1950).A studyof potassiumabsorptionby Jacobson, L., Overstreet, barleyroots. PlantPhysiol.25, 639-47. habit.I. Methodsofanalysisof Jefferies, R. L. & Willis,A. J. (1964).Studieson thecalcicole-calcifuge on fourspecies. J.Ecol. 52, 121-38. soil and planttissuesand someresultsofinvestigations kopcich. P,Viroda, v Breyovskych spolecenster Brno,29, k poynanirostlinnych Klika,J.(1936).Pfisp6vek 182-5. to thevegetaofthesoilanditssignificance ionconcentration Olsen,C. (1923).Studieson thehydrogen Ser. chim.15, 1-166. ofplants. C.R. Lab. Carlsberg, to thenaturaldistribution tion,especially solutions greenplantsinnutrient withhigher havingdifferent experiments Olsen,C. (1942).Waterculture Ser. chim.24, 69-98. ofcalcium. C.R. Lab. Carlsberg, concentrations ofchalkgrassland. J. Ecol. 48, 415-42. Perring, F. (1960).Climaticgradients Praeger,R. L. (1950).NaturalHistoryofIreland.London. of mineralnutrition on seedling problem.II. The effects Rorison,I. H. (1960).The calcicole-calcifuge growth in solutionculture. J. Ecol. 48, 679-88. responseto calciumwithinthespeciesFestuca A. D. (1961).Differential Snaydon, R. W. & Bradshaw, ovinaL. NewPhytol.60,219-34. IslandsandtheirVegetation. Cambridge. Tansley,A. G. (1949).TheBritish ofthecalcicoleandcalcifuge towardsan understanding Webb,D. A. & Hart,A. V. (1945).Contributions habitin someIrishplants. Sci. Proc.R. DublinSoc. N.S. 24, 19-28. R. L. (1959,for1958).The plantecologyoftheGordanovalley. Proc.Bristol Willis,A. J. & Jefferies, Nat. Soc. 29, 469-90. statusof thedunesoils. J. Willis,A. J. & Yemm,E. W. (1961).BrauntonBurrows:mineralnutrient Ecol. 49, 377-90. (Received9 April1964) R JE
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