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The Dependence of the Quantum Yield of Chlorella Photosynthesis on Wave Lenghth of Light Author(S): Robert Emerson and Charlton M

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The Dependence of the Quantum Yield of Chlorella Photosynthesis on Wave Lenghth of Light Author(S): Robert Emerson and Charlton M The Dependence of the Quantum Yield of Chlorella Photosynthesis on Wave Lenghth of Light Author(s): Robert Emerson and Charlton M. Lewis Source: American Journal of Botany, Vol. 30, No. 3 (Mar., 1943), pp. 165-178 Published by: Botanical Society of America, Inc. Stable URL: http://www.jstor.org/stable/2437236 Accessed: 24-06-2016 13:38 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://about.jstor.org/terms JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Botanical Society of America, Inc. is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany This content downloaded from 130.223.51.68 on Fri, 24 Jun 2016 13:38:58 UTC All use subject to http://about.jstor.org/terms THE DEPENDENCE OF THE QUANTUM YIELD OF CHLORELLA PHOTOSYNTHESIS ON WAVE LENGTH OF LIGHT 1 Robert Emerson and Charlton MI. Lewis MlEASUREMENTS OF the efficiency of plhotosyntihe- ble of photosyn-thesis. But owiing to the complexity sis in different wave lengths of light have receiitly of the plant's assimilatory mechaniism, it seems ap- been used as an approach to the problem of the propriate to inquire to what extent the activity of physiological role of the pigments accompanving chloroplhyll bears out expectation. chlorophyll. Dutton and M'ianning (1941) concluded Many investigators have accepted Warburg and from their measurements with the diatom Nitzschia Negelein's (1923) measurements of the efficiency closterium that the light absorbed by fucoxanthin of Chlorella photosynthesis in different parts of the and other carotenoid pigments was available for spectrum as satisfactory proof of the constancv of carbon dioxide assimilation. Emerson and Lewis the yield for light absorbed by chlorophyll. But (1942) made similar measurements with the blue these measurements were brought into question by green alga Chroococcus, and concluded that light the work of Manning, Stauffer, Duggar, and Dan- absorbed by the blue pigment phycocyanin gave a iels (1938) and Emerson and Lewis (1941) h-ave photosynthetic yield equal, within the experimental shown that Warburg and Negelein's results are error, to the yield for light absorbed by chlorophyll. subject to certain errors inherent in their method, They found, on the other hand, that the quantum and that their value for the quantum vield of photo- yield in the region of carotenoid absorption was synthesis should, therefore, be rejected. Since the much reduced, and regarded this as a clear indica- magnitude of the error arising from Warburg and tion that these pigments were not photosynthetic- Negelein's method may vary with wave length, their ally active with the full efficiency of chlorophyll and conclusions as to the dependence of quantum yield phycocyanin. However, when estimates of the pro- on wave length are also open to doubt. Further portion of light absorbed by the carotenoids were data on this point would have been desirable in any mnade from pigment extracts, and compared with case, because their conclusions were based only up- the dependence of quantum yield on wave length, on measurements in the yellow mercury line (578 evidence was obtained tlhat a fraction of the light m1u) and in a broad band of red light from 610 to absorbed by the carotenoids was probably available 690 m[u. Measurements were also made at 546 and for photosynthesis. The same technique has been 436 m[k, but at 436 the absorption was divided be- applied to the problem of carotenoid activity in the tween chlorophyll and carotenoid pigments, and at green alga Chlorella pyrenoidosa, and the results 546 the measurements were regarded as somewhat are reported in the present paper. uncertain because of the very dense cell suspen- In the work with both Nitzschia and Chroococ- sions required to absorb all the incident light in this cus, the quantum yield for light absorbed bv chlo- region where chlorophyll absorption is low.2 rophyll was measured in a wave length region The above considerations make evident the de- where all or nearly all of the total light absorbed sirability of more precise information as to the de- was absorbed by chlorophyll. This yield for light pendence of quantum yield on wave length, particu- absorbed by chlorophyll was then compared with larly in the red region, where chlorophyll is the the yield for spectral regions where light absorp- principal light-absorbing pigment. The absorption tion was divided between chlorophyll and accessory of light by the accessory pigments of Chlorella is pigments, to show whether or not the energy ab- restricted to a narrower range of the visible spec- sorbed by the accessory pigments was being used trum than is the case for Nitzschia or Chrobcoccus, for photosynthesis. In making this comparison it leaving a broader region for study of photosynthe- was assumed that the yield for the fraction of light sis in light absorbed by chlorophyll alone. If the absorbed by chlorophyll did not vary with wave yield for light absorbed by chlorophyll, and the length, but remained the same even when a large dependence of this yield on wave length, cani be fraction of the light was absorbed by other pig- established by measurements in a spectral region ments. The Einstein photochemnical equivalent law where chlorophyll alone absorbs all the light, this predicts that the primary photochemical action may imiake possible a more satisfactory analysis of should be proportional to the number of absorbed measurements in other spectral regions, where ab- quanta, so it inight reasonably be expected that the sorption is divided between chlorophyll and other quantum yield for light absorbed by chlorophyll 2 The dense cell suspensions made the respiration read- should be independent of wave length, at least over ings unduly large in proportion to the photosyntlhesis. The apparent necessity of using such dense suspensions the spectral range within which chlorophyll is capa- to achieve total absorption of the incident light may have 1 Received for publication October 6, 1942. arisen from the error associated with Warburg and The experimental work reported in this communication Negelein's method of measurement. When this error was was carried out during the years 1938-40 at the Carnegie avoided, we found no indications of incomplete light Institution of Washington Laboratory of Plant Biology, absorption, even with much thinner suspensions than they, Stanford University, Calif. used (cf. Emerson and Lewis, 1941, p. 801-802). [The Journal for Februarv (30: 83-163) was issued March 31, 1943.] AMERICAN JOURNAL OF BOTANY, VOL. 30, No. 3, MARCH, 1943. 165 This content downloaded from 130.223.51.68 on Fri, 24 Jun 2016 13:38:58 UTC All use subject to http://about.jstor.org/terms 166 AMERICAN JOURNAL OF BOTANY [Vol. 30, pigmiients of uncertain physiological activity. This compared with the absorption spectrum of the cells is the reason for the particular attention devoted to measured by direct methods. the red region in the present investigation. EXPERIMENTAL METHODS. MIost of tlhe teclh- In the course of the experimental work, two niques used have been fully described in earlier phenomena were encountered which raised special communications (Emerson and Lewis, 1939, 1941, problemiis in the measurement of the quantum yield 1942), and need only be mentioned briefly. Cells in certain portions of the spectrum. Exposure of were grown in the usual inorganic nutrient medium cells to the blue-green region sometimes caused a prepared in glass distilled water. Manganese, considerable inierease in the apparent rate of res- boron, zinc, copper, and molybdenum were added piration. An unexpectedly sharp decline in the in concentrationis of 0.5, 0.25, 0.025, 0.005, and quantum yield was observed in the far red. Sinice 0.05 parts per millioii respectively. Cultures were these two observations are probably at least as sig- grown in continuous incandescent illumination at nificant as the original purposes of the work, they a temperature of about 20?C. An iiioculum of about are discussed more fully than might appear to be 1 cmm. cells in 200 ml. of medium produced a necessary in view of their apparent treatmeiit mere- growth of about 300 cmm. of cells in from five to ly as difficulties of an experimental character. seveii days. Cultures of about this density were har- The same general plan has been followed as in vested for the photosynthesis measurements. The the correspondinig work on Chroococcus (Emerson cells were centrifuged out of the culture medium, and Lewis, 1912). To obtain evidence concerning washed twice in a mixture of 85 parts M/10 sodium the activity of accessory pigments from measuire- bicarbonate and 15 parts M/10 potassiumi carbon- ate, suspended in the same mixture, and pipetted x I l l l lI / ~ into the manonieter vessel. Twenty-five ml. of fluid containing about 300 cmin. cells was the usual filling for the manometer vessel. This concentration of cells was usually sufficient for total absorption 0 of the incident light, except for the green and in- frared regions of the spectrum, where denser sus- pensions were used. It was sometimes necessary to E 4 0 Red make a small correction for transmitted light, based C on visual estimation of the per cent transmitted. 0 Green The reliability of such estimates has been discussed A Blue in an earlier paper (1939, p. 819-20). Further ex- 2 / perience has shown that when the tranismitted light is less than 5 per cent, visual estimation introduces o no serious error.
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