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A Quantitative Study of Water-Culture Solutions A Quantitative Chemical and Physiological Study of Nutrient Solutions for Plant Cultures. by W. E.

Tottingham Review by: Walter Stiles Journal of Ecology, Vol. 2, No. 3 (Sep., 1914), pp. 182-185 Published by: British Ecological Society Stable URL: . Accessed: 08/08/2013 05:04
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Notices of Publications of General Bearing

the former method,till now based summations; hence forpracticalpurposesand forthe present, seems thus to be placed in closerlogical connection withthe solelyon phenologicalobservations, coefficients of chemical,physical and physiological processesthan has hitherto been temperature methodsof temperature betweenthe resultsderivedby these-two the case. (2) The similarity and roughly approximate;the ratiosof directsummation onlysuperficial integration is, however, season in the United States, for the mean frostless rangein magnitude, to efficiency summation climatic chart of 7-49 to a maximumof 10-44. (3) A rational and consistent from a minimum of these ratio values; on such a chart the marginal represents the geographicaldistribution characterised by low ratios,and the two main mountain are frequently regionsof the country systems(Appalachianand RockyMountains)appear to controlareas of high values. (4) There represents a climatic dimensionor seems to be no doubt that the ratio here broughtforward upon which appearsto be some sort of function of the dailynormaltemperatures characteristic, data within of the temperature has beenbased and of the timedistribution whichthiswhole study season. the periodof the mean frostless





"A quantitative chemical and physiological study of Tottingham, W. E. nutrient solutions for plant cultures." Physiol. Researches, 1, 1914, pp. 133-245. necessitate will be of greatuse to all whoseresearches solutions This paperon nutrient method;and or theuse ofthewater-culture ofplants solution intothenutritive investigations of on someaspects has muoh bearing it is onewhich an ecological paper notprimarily although ecology. ofthe andvaluable of a concise intothree parts. PartI consists r6siu?n Thepaperis divided thattheuse ofwater-culture It is shown work thathas beendoneup to nowonwater-culture. theideaof established about1860, ending period, periods.(1) The early three fallsinto methods from 1860 ofplants. (2) Themiddle period, material ofthenutrient as a source thesoilminerals and evolved classesof problems for certain thevalueofwater-culture 1900,established to about theresults bythe obtained for among solutions usein thismethod. Chief nutrient also standard for elements nutrition. Other oftheessential plant wasthedetermination in thisperiod method if the (0 3 0/0, or lower totalsalt concentration suchas an optimum factswerealso indicated, amount ofdifferent thatthe relative it was found although frequently), werechanged solutions uriouseffect. widelimitswithout inj any apparent producing rather salts couldvarywithin and saltswerealso indicated.(3) In the third of certain and excretion absorption Selective saltsthatis indicated, thancomplete of ionsrather absorption it is the selective period present altered. Theimportance becomes solution ofthenutrient thecomposition and so bythismeans has also been solution in thenutritive thevarious elements balancebetween of a physiological recent years. during emphasised in relative out the great variations finishes thispartof his paperby pointing The author that andso concludes solutions, nutrient standard in thevarious ofdifferent substances amounts of proportions valuesofdifferent oftherelative tomake" a thorough investigation it is justifiable thebeginning constitutes ofthepaper media." Theremainder in suchculture thesaltsemployed ofsucha pieceofwork. as a resultof whichthe most is made of Knop's solution, study In PartII a chemical to make ofcourse is described.It is desirable solution ofmaking method up a stock satisfactory withstability.Tottingham as is consistent whichis of as higha concentration up a solution theother nitrate one of calcium containing alone,theother twostock solutions, using suggests of 23?C. in solution is stableforabout14 daysat a temperature salts. The latter necessary as K2HPO4 or KH2PO4 is used as the sourceof of 9 ?/. or 100/0 according a concontration

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A QuantitativeStudy of Water-culture Solutions


phosphorus. The precipitate above the limitof stability is almostentirely calcium sulphate,not calcium phosphateas has been supposed. In Part III of the paper we have a recordof the experiments which werecarriedout with a view to determining the most suitable nutrientsolution. Young wheat plants were used, and were grownforapproximately thirdof the fullgrowth the first periodof wheat. Colourlessflint glass bottlesof 250 c.c. capacitywere used, and six plants were grownin each bottle. They remained24 days in the nutrient solutions,whichwere changedevery thirdday. At the end of the growth periodtheyweredriedand the dryweightsof shootsand rootstakenseparately. Preliminary series of experiments were made in which Knop's solutionwas used in various concentrations but in whicha constant ratiobetween the variousconstituent salts was maintained throughout. In the case of each solutionits osmoticpressureis calculated fromelectrolytic dissociation data, but as no accountis taken of the influence on the dissociationof one another produced by severalsalts together in the same solution,the numbers forthe osmoticpressure are onlyapproximate. Tottingham'sresultsshow that concentration has practically no influence on the amount of drymatter producedover a veryconsiderablerange,namelyfrom a concentration of total salts of 001 about one of 1-4per cent. It is concludedalso thatmonopotassium phosphateis a better salt to use as a sourceof phosphorus than the dihydrogen phosphate, and it is the former substancethat was used in the subsequentseries. seriesthe In the further experiments threemain seriesof cultureswerearranged. In the first total concentration of salts was 06 0/0, whichlies well withinthe range of concentrations giving optimum growth. In thesecondseriesthe concentration of salts was 0-010/0. This concentration " was selectedas having been shown in the preliminary culturesto lie considerably belowthe similar to that of manynaturalsoil optimumi," and also because the concentration is probably solutions. In the thirdseries the concentration, 20 0?/, was above the limits of optimum concentration, and also presented conditionsof concentration similarto thosewhich obtain in alkali soils. In each of these series a numberof nutrient solutionswere obtainedbyvarying the relative proportionsof the four salts Ca(NO3)2,KNO3, MgSO4, KH2PO4, but within each series the osmoticpressuresof the various solutions remainedapproximately constant. The dryweights weretakenand the resultsplotted on the triangle diagramarrangement used in physicalchemistry and introduced intobotanicalconsiderations by Schreiner and Skinner2* The chiefresultsof these experiments are as follows: of thenutrient 1. Wherethetotalconcentration solutionwas 0 6?/.. (a) Wherethe osmotic MgSO4 ratio of was greater than 2-0 (=a molecularratio of 3 23), a verymarkedand curious Ca(N03)2 injuryto the plants was observed. The tips of the leaves "rolled backwardlaterallyto form " and thiswas generally a stiff needle-like structure and ultimatebleaching preceded byyellowing of the tip of the leaf3. Later the youngleaves became limp and spirallycoiled in the middle or No injury was observedwhen the ratio MgSO4 was below 0-25 (= a Ca(N03)2 wsblw05( molecularratio of 0 40). The extentof the injuryin cultureswherethis ratio was intermediate towards the base. 1 However,the actual difference in dryweightbetweenthe culturein this concentration and is so slightthat the difference those in the optimum concentrations may quite possiblylie within the ralnge of experimental error. 2 "Some effects of a harmful organic soil constituent." U.S. Dept. of Agric.Butr. of Soils, Bull. 53, 1910; also, " Ratio of phosphate,nitrateand potassiumon absorption and growth." Bot. Gaz., 50, 1910, pp. 1-30. 3 This formof injury was observedby the reviewerin water-cultures of rye grownin the was as high as 14, while it was only springof 1913. In this case the molecularratio Mag(N04) low concentrations much lowerthan in Tottingham's experiments (0 6). In very ( 01) Tottingham non-existent. foundthe injurypractically
slightly in evidence when the ratio was



2-8. The total concentration (075) was however

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Notices of Publications of General Bearing

between0-25 and 2-0, was presumaably determined by the concentration of the othersalts in the medium. (b) The best growthof shootswas obtainedin solutionshavingthe salts in approximately the followingosmotic proportions: 2-5 KH2PO4: 2-5 MgSO4: 4 0 CaNO3 1 0 KNO:3 to molecular proportionsof 2-66 : 2-97: 2-9 : 1 00, a considerably corresponding different from arrangement Knop's. The best solutionforthe growthof rootswas howeverfoundto be quite different, namelyone in whichthe osmoticproportions were1 00: 0-25 : 0-25 : 1 00 (correto molecularproportions Fponding 1-07: 0 30: 0-18: 1 00)1. (c) The development of the shoots was not proportional to thatof the roots. Livingston had foundthis to be so in the case of wheat growingin soil cultures2,so Tottinghamsupposes that while in a liquid hnedium " the conof the variousmolecules and ions are not greatly centrations influenced of the by the metabolism plant,and considerable differences in rootarea mayhave slightinfluence uponthe growth of tops, on the otherhand,theactivitiesof theplant may(through in soil cultures, of carbon-diexcretion of the soil solutionin variousirons,and the extentofthe root oxide, etc.) alterthe concentration system maythenbecomea much moreimportant factor in the growth oftops." This explanation does not seem to be convincing. 2. It the cultur es wherethetotal concentratioin of the solution was 0-010/0. (a) Both shoots and rootsoftheplantwereelongated and slender, thisbeingleast marked in the cases ofrootsgrowing in solutions lowestin concentration of MgSO4and highestin KNO3concentration.(b) In some culturesa yellowing and dyingof the basal leaves occurred, probably due to deficiency of nitrate. The low yields in some cases appear to be due to the same cause. (c) The peculiarmagnesium wherethe ratioof M(g(04) cultures injurywas absent,but the greatest yieldswereobtainedfrom was low, and the least yieldsfrom cultures wherethe ratiowas high. (d) The bestproportion of salts for the growthof shoots was consideredto be in osmoticproportions, 0 5 KH2PO4: 2-0 MgSO4 : 15 Ca(N03)2 : 10 KNO3 (corresponding to molecularproportions of 0 34: 2-00 : 100: 1.00). In the case of roots the best osmoticproportions were06 : 02 : 0 2 : 10 (colresponding of 0-41 : 0-20 : 0-13 : 1 00). (e) As in the preceding to molecularproportions seriesthe yieldof shootsboreno relationto the yieldof roots. 3. In thecultures where thetotalconcentration of salts was 2 0 0/0. (a) Root development was affected the series,but chiefly adversely throughout wheretherewas an excess of any one of the nutrient salts. An excess of hydrogen, calcium or magnesium ions appearedto be mostinjurious. (b) A high proportion of calciumappearedto cause wilting of the leaves and ultimately the death of the plant. (c) In some culturesthereappeared a yellowing of the basal leaves followedby death. This mighthave been due to a low proportion of magnesium sulphate,butwas not clearly so. (d) In otherculturesthe tissuebetween the leaf veins becamebleached. This was probably caused by the high concentration of salts. combined with certain unfavourable proportions (e) Magnesiuminjuryoccurredas in the firstseries but no simplerelationbetweenmagnesium and calcium could be obtainedin this case. It is clear that the antagonism between calciumand magnesiumis much influenced of salts for shoot by concentration. (f) The best proportion was found to be *22 KH2PO4 : *67 MgSO4 : 33 Ca(NO3)2: 100 KNO3 in osmotic development to molecularproportions proportions (corresponding 0-23: 081 : 0-25: 1 00). In the case of roots the best osmoticproportions was consideredto be 04 : 1-8 : 0-8: 10 (corresponding to of 0-41 : 2-20 : 0 59 : 1-0)3. (g) There was not much correlation molecularproportions between growthof roots and shoots in this series. (h) A great decrease in the ratio of waterloss by to the yieldof shoot accompaniedexcessivemagnesiuminjury. In the first series transpiration no such correlation was obvious,and the decrease in waterloss in this series is presumably due to the reducedleaf surfacein the much injuredplants. The transpiration per unit of dryweight 1 These appear to be the correct in the summary numbers, and are so givenby Tottingham of the paper. Those actuallygivenat this stage of his paperare quite different. 2 " Note on the relationbetween the growth of rootsand of tops in wheat." Bot. Gaz. 41, 1906, pp. 139-143. 1 The numbersgivenby Tottingham at this stage ofhis paperare quite different.The ones his summary the correct quotedhere are takenfrom and are presumably ones.

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A QuantitativeStudy of Water-culture Solutions


was also much less than in the second series,a resultwhich was also found,thoughto a much of nutrient solution. less degree,in the cultures in weakestconcentration to call attention In conclusionit wouldseem necessary to two criticisms whichmay be made on the workhere dealt with. The first criticism deals withthe quantity of solutionin which the together cultures weregrowing. In nearlyall cases each cultureconsistedof six plants growing in one culturejar containing 250 c.c. of solution. This means only42 c.c. of solutionper plant, and althoughthe solutions werechangedevery threedays and althoughthe plantsweresmall, yet it is quite possiblethat the plants wereabsorbingsalts quite fast enoughto alter appreciably the of solution. proportion of the varioussalts in such a small quantity of plants growing in water In the secondplace, no account is taken of the greatvariability six plantswas grown culturesunderexactlythe same conditions. As onlyone culturecontaining in the same solution it is verypossible that some of the differences of dry weighton which due to this. But in the absence of any data in emphasisis laid, are withinthe limitsof the error this is so. regardto probableerror it is not possibleto say whether But regarded generally it would appear that Mr Tottingham'sworkhas been most carefully justified. It is probably difficult forthose carriedout, and most of his conclusionsare probably who have not conductedwork involvingthe use of water-cultures to realise the great amount of labour involvedin carrying to a successfulconclusionsuch a piece of workas that here consideredwiththe care thathas evidently been bestowed upon it. To the ecologistof coursethe interestof this worklies verylargelyin the applicationof the to the soil solution. How farit is permissible to arguefrom resultsobtainedto questionsrelating in water-culture to problemsof the soil it is impossibleat presentto say. That effects produced is obvious,but we mustwait before such effects musthave some bearingon soil problems we can differences in concentration telldefinitely to whatextent of the soil solutionor in theratiobetween of different substancesin the soil solution are really operativein influencing the distribution vegetation.


RECENT Snell, K.






deutsch. bot. Ges., 30, 1912, pp. 361-362.

Transpirationsstromder Wasserpflanzen." Ber. d.

(III) (IV) (V) (VI)

Solereder, H. "Systematisch-anatomische Untersuchungendes Blattes der Hydrocharitaceen." Beih. z. Botan. Centralbl., 30, Abt. 1, 1913, pp. 24104, 53 figures. Beyrer, H. "Beobachtungen iiber das Etiolment bei Wasserpflanzen." in [etschen,14, 1913, pp. 3--16, 1 plate. Jahresber. Staats.-Gymn. D. T. " The determinative action of environic factors MacDougal, upon Neobeckia aquatica Green." Flora, 106, pp. 264-280, 14 text-figures. Esenbeck, E. "Beitrage zur Biologie der GattungenPotaintogeton unid Scirpus." Flora, 107, 1914, pp. 151-202, 59 text-figures. Gertz, A. "Pildammarnas vegetationeftertorrlaggningen." Bot. Notiser, 1913, pp. 113-130.

The papers cited above are selected,fornotice here, from the veryextensiverecentliterature on the ecologyof freshwater as a supplementto the collective aquatic and marsh vegetation, notice of papers on the subjectwhich appeared in an earliernumberof this JOURNAL (1, 1913, is growing so rapidlythat we have again omitted pp. 107-115). This literature noticesof those contributions which appear to be of minor general interest to the ecologist, and have as a rule

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