historical aspects of ultraviolet action spectroscopy*

6
Photochemistry and Photobiology, 1997, 65s: 123s-128s Historical Review Historical Aspects of Ultraviolet Action Spectroscopy* Thomas P. Coohillt Ultraviolet Consultants, Bowling Green, KY, USA ABSTRACT One of the fundamental methods for analysis of an or- ganism’s responses to UV radiation is the construction of the wavelength dependence for a given bioeffect, termed an action spectrum. At minimum, this demonstrates the organism’s response to different spectral regions; in some cases, it points to the target chromophore for the effect. The historical importance of certain action spectra and the development of refined techniques for their con- struction is the subject of this review. Wherever possible, I have kept to the original spellings, terms and identifi- cation of wavelengths. This includes using angstroms and nanometers (but not millimicrons) as the authors did. I preferred to reference papers with the most complete set of data rather than the earliest reports. INTRODUCTION The trouble with history is there’s no end to it. Accordingly, I have selected a small number of studies for this review. I apologize to those authors, living and dead, who warranted inclusion but were not cited. In 1836, 109 years after the death of Newton, Daubeny (1) reported a study that proposed to, “test . . . whether the several solar rays act upon plants with equal or with different degrees of energy . . . (and whether) the effect of light upon plants corresponds with its illuminating, rather than with its chemical, or its calorific influence . . . .He quotes the work of Senebier (2) who said, “the green color of leaves . . . is produced most rapidly under the action of the violet ray . . . .” Using a variety of glass and liquid (including port wine) filters, he found, “the most luminous rays were most influ- ential . . . (i.e.) orange . . . more than red or green . . . yet in some instances blue and purple . . . more . . . than orange . . . .Hardly the basis for an action spectrum, but a begin- ning. By the end of the century, Englemann (3) and Draper (4), produced what may be termed the first crude visible action spectra for the production of chlorophyll, pointing to *Presented at the American Society for Photobiology in St. Louis, MO, July 1997, as part of the Landmarks in Photobiology Sym- posium celebrating the 25th Anniversary of the founding of the Society. tTo whom correspondence should be addressed at: Ultraviolet Con- sultants, 652 East 14th Street, Bowling Green, KY 42101, USA. Fax: 502-782- I35 I. 0 1997 American Society for Photobiology 003 1-8655/97 $5.00+0.00 it as the molecule most responsible for plant growth and setting in motion the idea of the practicality of such studies. The Draper paper (4) is particularly interesting to read for its rip-roaring condemnation of “Mr. Hunt from Falmouth (England),” who disagreed with results Draper published 48 years earlier! We were on our way. SUN “BURN” AND SKIN CANCER In the 18th century, Senebier (2) stated “peasants working much in the open have paler skin in covered areas than ex- posed areas, and, if exposed for years to sunlight, the face and hands appeared thickened and tanned, while in contrast, the covered areas retain their white appearance.” This was an early observation of the effects of sunlight on human skin. Near the end of the second decade of the present century, the German physicist Wilhelm Hausser contracted tubercu- losis. Sent to the mountains for the air, he noted “a long hike on a glacier, in the afternoon hours under a burning sun had almost no effect, while . . . a brief sojourn on snow at noontime resulted in a severe sunburn.” He wondered which of the “ultraviolet component(s) of the radiation” was caus- ing this effect and began a series of elegant experiments to find out. He got around the obvious constraints of working with humans by establishing a simple assay. The inner fore- arms of patients were exposed to the individual lines of a mercury lamp spectrum, separated by two quartz prisms. This action spectrum for human erythema (5) had a sharp peak at 297 nm. This paper also seems to be the first use of the term “action spectrum,” and the spectrum itself is the first of the “modern” type. Hausser’s results would be con- firmed by others. In 1967, Berger e? al. (6) reexamined the action spectrum data of Hausser and Vahle (5) and reported additional studies. They were able to eliminate errors due to “stray light” and account for the discrepancies in effect at short wavelengths noted by previous workers, by noting the differences in the time of evaluation of the skin response and to the degree of “redness” chosen to establish this min- imal erythema! dose (MED).$ They agreed with Hausser in all major respects. Those who do not read German might find a starting point to follow the progress of action spectroscopy in human skin in a paper by Coblentz et al. (7). He reports his own work, $Abbrrviutions: AAS, analytical action spectrum; AS, action spec- trum; MED, minimal erythemal dose; SCUP-h, skin cancer Utrecht-Philadelphia-human; TSE, transmissible spongiform en- cephalopathies. 123s

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Page 1: Historical Aspects of Ultraviolet Action Spectroscopy*

Photochemistry and Photobiology, 1997, 65s: 123s-128s

Historical Review

Historical Aspects of Ultraviolet Action Spectroscopy*

Thomas P. Coohillt Ultraviolet Consultants, Bowling Green, KY, USA

ABSTRACT One of the fundamental methods for analysis of an or- ganism’s responses to UV radiation is the construction of the wavelength dependence for a given bioeffect, termed an action spectrum. At minimum, this demonstrates the organism’s response to different spectral regions; in some cases, it points to the target chromophore for the effect. The historical importance of certain action spectra and the development of refined techniques for their con- struction is the subject of this review. Wherever possible, I have kept to the original spellings, terms and identifi- cation of wavelengths. This includes using angstroms and nanometers (but not millimicrons) as the authors did. I preferred to reference papers with the most complete set of data rather than the earliest reports.

INTRODUCTION The trouble with history is there’s no end to it. Accordingly, I have selected a small number of studies for this review. I apologize to those authors, living and dead, who warranted inclusion but were not cited.

In 1836, 109 years after the death of Newton, Daubeny (1) reported a study that proposed to, “test . . . whether the several solar rays act upon plants with equal or with different degrees of energy . . . (and whether) the effect of light upon plants corresponds with its illuminating, rather than with its chemical, or its calorific influence . . . .” He quotes the work of Senebier (2) who said, “the green color of leaves . . . is produced most rapidly under the action of the violet ray . . . .” Using a variety of glass and liquid (including port wine) filters, he found, “the most luminous rays were most influ- ential . . . ( i .e . ) orange . . . more than red or green . . . yet in some instances blue and purple . . . more . . . than orange . . . .” Hardly the basis for an action spectrum, but a begin- ning. By the end of the century, Englemann (3) and Draper (4), produced what may be termed the first crude visible action spectra for the production of chlorophyll, pointing to

*Presented at the American Society for Photobiology in St. Louis, MO, July 1997, as part of the Landmarks in Photobiology Sym- posium celebrating the 25th Anniversary of the founding of the Society.

tTo whom correspondence should be addressed at: Ultraviolet Con- sultants, 652 East 14th Street, Bowling Green, KY 42101, USA. Fax: 502-782- I35 I .

0 1997 American Society for Photobiology 003 1-8655/97 $5.00+0.00

it as the molecule most responsible for plant growth and setting in motion the idea of the practicality of such studies. The Draper paper (4) is particularly interesting to read for its rip-roaring condemnation of “Mr. Hunt from Falmouth (England),” who disagreed with results Draper published 48 years earlier! We were on our way.

SUN “BURN” AND SKIN CANCER In the 18th century, Senebier (2) stated “peasants working much in the open have paler skin in covered areas than ex- posed areas, and, if exposed for years to sunlight, the face and hands appeared thickened and tanned, while in contrast, the covered areas retain their white appearance.” This was an early observation of the effects of sunlight on human skin. Near the end of the second decade of the present century, the German physicist Wilhelm Hausser contracted tubercu- losis. Sent to the mountains for the air, he noted “a long hike on a glacier, in the afternoon hours under a burning sun had almost no effect, while . . . a brief sojourn on snow at noontime resulted in a severe sunburn.” He wondered which of the “ultraviolet component(s) of the radiation” was caus- ing this effect and began a series of elegant experiments to find out. He got around the obvious constraints of working with humans by establishing a simple assay. The inner fore- arms of patients were exposed to the individual lines of a mercury lamp spectrum, separated by two quartz prisms. This action spectrum for human erythema ( 5 ) had a sharp peak at 297 nm. This paper also seems to be the first use of the term “action spectrum,” and the spectrum itself is the first of the “modern” type. Hausser’s results would be con- firmed by others. In 1967, Berger e? al. (6) reexamined the action spectrum data of Hausser and Vahle ( 5 ) and reported additional studies. They were able to eliminate errors due to “stray light” and account for the discrepancies in effect at short wavelengths noted by previous workers, by noting the differences in the time of evaluation of the skin response and to the degree of “redness” chosen to establish this min- imal erythema! dose (MED).$ They agreed with Hausser in all major respects.

Those who do not read German might find a starting point to follow the progress of action spectroscopy in human skin in a paper by Coblentz et al. (7). He reports his own work,

$Abbrrviutions: AAS, analytical action spectrum; AS, action spec- trum; MED, minimal erythemal dose; SCUP-h, skin cancer Utrecht-Philadelphia-human; TSE, transmissible spongiform en- cephalopathies.

123s

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and compares and analyzes the work of Hausser and Vahle (S), and Luckiesh (8). All three studies found a sharp peak at 297 nm, a deep trough at 280 and 313 nm and a general increase in effect for wavelengths as short as 248 nm (al- though they disagree in magnitude of effect at the shorter wavelengths). Both Luckiesh (8) and Coblentz et al. (7) showed that reciprocity held. In all cases they used the pa- tient as an indicator and estimated the dosage by the MED. In 1927, Hausser and Vahle (9) published a comprehensive review of their work, adding important points, e.g. the ab- sorption properties of the skin. They had the luxury of start- ing a scientific paper “With the change of seasons, with the blossoming of plantlife in the spring and its death in the fall, there is also seen a peculiar and note-worthy alteration in the appearance of, particularly nordic, people.” Fred Ur- bach’s translation of this work is worth reading (10).

An action spectrum for photocarcinogenesis, the most widespread of all human cancers, proved more difficult to construct and required the use of animal models (mainly mice). The association of solar exposure and skin cancer dates back over 100 years, and Blum ( l l ) , using cut-off filters, suggested the target could be nucleic acid. Even so, Forbes et al. (12). as late as 1978, stated “virtually no direct data are available on dose-response or action spectra for human skin UV photocarcinogenesis; furthermore, such data are not likely to come into existence soon.” Lack of progress was partly due to the general assumption that the action spectrum for human carcinogenesis would follow that for erythema. In fact, this appears to be the case. De Gruijl and Van der Leun have recently (13) produced a part analytical and part experimental action spectrum for human (and one for mice) photocarcinogenesis. Called the skin cancer Utrecht-Philadelphia-human (SCUP-h) action spectrum, it denotes the two laboratories most responsible for the data, much of it from experiments with hairless mice. The SCUP-h combines what is known from mice and human data and estimates of skin transmission, etc. The peak response is at 299 nm, with a rapid lowering of effect at both longer and shorter wavelengths (about a factor of 100 below the peak value at 270 nm and loo0 below the peak at 330 nm). The long wavelength data follow, within reason, the theo- retical analysis of Setlow (14).

A recent, and to some surprising, action spectrum for ma- lignant melanoma is that of Setlow, et ul. (15). Realizing the difficulty in obtaining such a spectrum for a human response, he utilized a platyfish model that developed melanomas in a matter of months. This spectrum followed the normal DNA- related curve for mammalian cell death and mutagenicity for wavelengths in the W B . However, in the UVA and violet regions of the spectrum, the response was much higher than either cellular response. Thus, unlike erythema and nonme- lanoma skin cancer, malignant melanoma is easily elicited by UVA and even visible radiation.

FIRST ACTION SPECTRUM FOR CELL DEATH The importance of bacteria. During much of the first half of this century, it was generally believed that proteins were the genetic material. An example of this was the work of Harris and Hoyt in 1917 (16) that showed that irradiation of para-

mecia through a solution of amino acids greatly lowered cell killing. They concluded “Our results are therefore decidedly in harmony with the view that the susceptibility of proto- plasm (here paramicia) to ultra-violet light is conditioned by the selective absorption of the toxic rays by the aromatic amino-acids of the proteins.” Contradicting this view, and of seminal importance to molecular biology, was the set of action spectra for bacterial cell death from the laboratory of Frederick L. Gates at the Rockefeller Institute (17-21). In the 1920s, he irradiated a variety of cells and viruses, most notably the bacteria Staphylococcus aureus and Escherichia coli. One of the requirements for construction of an analyt- ical action spectrum ( U S ) is that the target chromophores see most of the incident beam. That is, absorption and scat- tering should be minimal. This is almost never the case for the W C in plants or animal cells, due to their thickness and pigmentation. These organisms are discussed below, but it should be noted here that Gates’ work with bacteria pro- duced the first A A S and pointed to the identification of the genetic material. He measured, among other things, the ab- sorption of a monolayer of bacteria (17) and found it “so transparent that objects may be seen through it clearly and without distortion . . . ,” thus meeting the transmission re- quirement of an AAS.

In 1928, Gates (18) gave the first brief description of what was to come out of his laboratory. It disputed the conclu- sions of Harris and Hoyt (16). Gates stated that “while the relation of thymonucleic acid to cell growth and reproduc- tion remains a matter of conjecture . . . the reciprocal of the batericidal curve matches the absorption curves for cytosine, thymine, and uracil more closely than . . . those for various amino acids suggested by Harris and Hoyt.” This will prove to be his strongest statement on the matter. He ends this paper by suggesting that his observations “seem worthy of note, without further comment.” This is followed by a series of papers in the Journal of General Physiology that begin in 1929 (where he notes his experimental procedures and de- scribes the shape of the UV survival curves (19)); also in 1929 (where he tries to establish the wavelength boundaries for cell killing (20)) and, his 1930 publication (17) that is considered by some to be the most crucial action spectrum ever published. Of historical interest is Gates’ publication (21) where two complete action spectrum for cell death of the same shape are shown, one from 1923 and one from 1927. Comparison of these spectra for S. aureus shows that the spectrum in the 1930 paper is from the 1923 data. These spectra paved the way for a host of studies by molecular biologists.

Figures 2 and 3 of Gates (17), plot the reciprocal of the energy required to produce cell killing in two bacteria. The peaks for each curve are at 265 nm. There were no data for any other wavelength between 254 nm and 280 nm, probably due to the low mercury lamp output at 258, 260, 270 and 275 nm. Based on the shape of the spectrum, Gates states “the simple conclusion of former investigators that the shorter the wavelength of ultraviolet light the greater the bactericidal action is in error.” He equivocates more here about the nature of the target material than he did in 1928 (18). ending the paper with “these curves . . . point the way in a future search for the specific substance, or substances, involved in the lethal reaction.” He claims that “A final

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paper in this series will discuss structural and chemical units of bacterial protoplasm that may prove to be involved in the reaction which results in the organism’s death.” Alas, this promised work (to be paper number IV in the series) was never published.

Gates died in 1933. Although some of his work was pub- lished by others posthumously, none contained the promised identification of the “chemical units . . . involved . . . in the organisms death” (17). Why this is so is a puzzle. Perhaps the continued belief among biologists that protein was the genetic material made Gates’ followers hesitant.

Ehrismann and Noethling (22) published a study of cell death as a function of UV wavelength for a variety of mi- croorganisms. Their results agreed with Gates.

In the late 1930s and early 1940s, a series of action spec- tra (AS) were produced that measured both lethality and mu- tation (see below) and agreed with the concept that nucleic acid was the genetic material. A comprehensive review up to 1930, which includes a discussion of techniques and tar- gets, is contained in Weinstein (23). Things were getting clearer .

HOLLAENDER AND OTHERS

Other authors adopted the methods of Gates. Central among them was Hollaender and his associates who published a series of papers. Hollaender and Claw (24) provide a thor- ough numerical analysis for survival curves of E. coli after exposure to UV and conclude, “(the) mechanism by which ultraviolet radiation inactivates bacterial cells is a question under considerable discussion . . . one fact stands out . . . the sensitivity curve . . . has a maximum at 2650A” (Note here and in what follows, that Hollaender, like Gates, does not utilize the mercury line at 2602A.) He provides action spec- tra that peak at 2650A for both killing and mutation pro- duction in the fungus Trichophyton mentagrophytes (25-27). Hollaender and Emmons (25) has the most complete data, but their previous work (26) shows in fig. 3 that only when the cells are irradiated, so as to allow some UV to penetrate the cell completely, does the spectrum have a smooth peak like in bacteria. This is an important example of the trans- mission constraint for AAS. Although it could be argued that cell death may not be directly related to the genetic material, a similar argument would be less tenable for mutation. Yet, even after these studies, they include (27) the caution “it is probably somewhat dangerous to overemphasize the impor- tance of nucleic acid , . . (it may be) only the ”absorbent“ agent, (which) then transfers . . . (the) energy to . . . pro- tein.” It is worthy of note that this paper (27). which ends with a discussion among several experts, contains not one question dealing with whether nucleic acids are the target material or the genetic agent.

By 1944, Hollaender and Oliphant (28), working now with viruses known to have high nucleic acid content, state, “it is quite possible that the high sensitivity . . . at about 2600A is based on the important function desoxyribose nu- cleic acid plays in biological activities.” By 1954 (29), au- thors refer to “action spectra of the nucleic acid type.” Ac- tion spectroscopy had indeed arrived.

There are many reviews covering action spectra in micro- organisms. An early comprehensive series of reviews on the

broader questions of organism response to UV are contained in Vol. I1 of the series Radiation Biology (30) edited by Hollaender. The chapters on genes and chromosomes, pro- tozoa and invertebrate eggs. viruses, bacteria and fungi, are particularly germane and include most of what was known until then about UV effects on microorganisms and model systems (e.g. fruit flies). The long review by Loofbourow (31) describes much of the mathematical analysis necessary for AS studies; Giese (32) produces a useful chart of AS up to 1944 that includes a column entitled “possible absorbing substances.’ ’

ACTION SPECTRA FOR PLANTS

The first studies that might be considered to be the begin- nings of action spectroscopy involved plants and were men- tioned above. But a substantial problem in attempting such work with plants is the presence of numerous pigments that can shield target molecules. Transmission of UV is often so restricted that the results obtained may not point to a target chromophore. Usually, sophisticated analysis is required and, even then, only vague answers possible. Hausser and Oehmcke (33) avoided these problems by irradiating banana skin with the spectrally separated output from a mercury lamp and noting pigment darkening on the surface. The re- sults were clear lines corresponding to the mercury spec- trum, arranged in sequence. It is interesting to note their report that the wavelengths 366, 405 and 436 nm interfered with the formation of pigment by UVB. Were they the first to discover photoreactivation?

Stadler and Uber, presenting data collected earlier (34), produced an action spectrum for genetic effects in maize. It closely followed the absorption spectrum of thymonucleic acid (which was plotted alongside their experimental points). This type of presentation is now common, although the com- parison curve is usually DNA. In addition, they included a brief review of other mutational action spectra, useful be- cause they summarized German work. They cite the data of Noethling and Stubbe (35) who exposed pollen of Antirrhi- num majus and found mutations well into the UV, although their conclusions were hindered by the appreciable absorp- tion of their pollen suspension. However, Knapp and co- workers, (36) using the sperm of S. donnellii (which is less than 1 Fm thick and much more transparent), found 2650A to be the most effective wavelength for mutation production.

Florence Meier in 1936 (37) presented a wonderfully comphrehensive paper on the lethal effects of UV on Chlo- rella. The first to utilize a large number of wavelengths (20 in the range 2250-3022A), she also included the mercury lines 2537, 2602, 2652, 2699 and 2753A that bracket the nucleic acid peak. She showed that 2602A was the most effective wavelength for cell killing and that reciprocity of time and dose held over a factor of 12. This paper is re- markable for the detail it presents and the large number of studies she completed (some of the data are from her earlier work in that decade). She does not speculate on the nature of the target molecule.

Phycomyces, with its easily observed “growth-light” re- sponse has been a favored organism for investigators. Castle (38), published an action spectrum for this effect that ex- tended from the visible into the long wavelength UV. Del-

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126s Thomas Coohill

bruck and Shropshire (39) and Curry and Gruen (40) ex- tended this work to shorter wavelengths and demonstrated several additional peaks in the UV. Galland (41) reviewed the action spectra for this species, including some mutants, in 1983.

A large number of plant AS, many of which are men- tioned in the reviews cited below, concerned growth and photosynthesis. Because of the difficulties of structure and pigmentation in plants, a more recent AS using spinach thy- lakoids can be cited that suggests that photosystem I1 is more sensitive to UV than photosystem I(42). Reviews for action spectra in algae (43) and higher plants (44,45) are available.

Of current interest are AS produced from field data (46). These usually do not meet the rigorous criteria for AAS but are important for understanding the reaction of organisms in nature. A brief review of such aquatic spectra is found in Booth et al. (47).

MAMMALIAN CELLS

Shortly after the publications of Gates, attempts were made to determine the agent that would cause UV killing in mam- malian tissue. As if to underscore the difficulty of working with material that absorbs substantially in the UV, Mayer and Schreiber (48) completed an AS study similar to that of Gates but utilized hanging drop cultures of mammalian tis- sue. They found that growth and morphological change could be increasingly inhibited by UV wavelengths down to about 270 nm. However, their curve flattened out at shorter wavelengths due to the opacity of tissue in this range. After a long hiatus, the advent of single cell tissue culture tech- niques allowed further work to proceed. In 1965, Chu (49) published an AS for chromosomal aberrations in hamster cells that gave a flat response between 240 nm and 280 nm. He also showed a marked increase in effect if the thymine analog 5-bromodeoxyuridine was present in the cells, which was consistent with the view that DNA was at least partially responsible for these effects. In 1968, Todd et al. (50) re- ported the first AS for mammalian cell killing. It was shifted about 50 nm to the long wavelength side from the spectrum of Gates. Further work showed that this shift could be ac- counted for by cytoplasmic absorption and that the spectrum followed the production of pyrimidine dimers in these cells (51). No differences were found for action spectra in any human photosensitive cell line studied (52,53), That is, DNA was the primary target. A review current to 1984 is con- tained in Coohill (51).

VIRUSES

The first AS for a virus was the one of Rivers and Gates in 1928 (54). They showed a peak for vaccine virus sensitivity at 2652A. This was followed by a series of articles that showed the same wavelength sensitivity for the bacterio- phage of S. aureus (21), influenza virus (28,55), the T1 bac- teriophage of E. coli (56), viral lysis in E. coli (57), herpes simplex (58) and potato virus X (59). Many commented on the fact that the same absorbing compound that inactivated cells, might be inactivating viruses. The latter studies as- sumed this was a nucleic acid.

PRIONS AND AS In 1732 a clinical description of the disease scrapie noted the unbearable itchiness that caused sheep to rub against fences or trees. This was the first of the transmissible spon- gifonn encephalopathies (TSE) that affect many animals, most recently the “mad cows” of Britain. The human dis- ease Creutzfeldt-Jakob was linked to these TSE after kuru was discovered in the Fore people of New Guinea. It was transmissible, in the case of the Fore, by cannibalism. It was thought to be a virus, perhaps a “slow virus.” But there was something odd about this virus. You could subject it to high doses of gamma rays, disinfect it with formalin or treat it with UV, but you could not kill it. What type of nucleic acid was this?

In 1967, Alper et al. (60) irradiated the scrapie agent with much higher fluences of UV. Their study charted the effec- tive killing of Micrococcus radiodurans at UV fluences far below those that just began to show some dimunition of infectivity by scrapie. By then the pendulum started by Gates, which identified nucleic acid as the replicative ma- terial, had swung far enough that Alper could state, “The responsibility of the nucleic acids for replication and genetic control is so firmly established . . . that the agent (scrapie) must be devoid of nucleic acid . . .. and might be a repli- cating polysaccaride.” Lataqet et al. (61) managed to in- activate scrapie with enormous fluences of UV and produce an AS that he stated was “other than a part of a nucleic acid molecule or of a nucleoprotein complex.”

Current-day prion theory supports some of these early conclusions. The infectious agent may be a protein or a small amount of nucleic acid somehow protected by a protein. Ac- tion spectroscopy pointed the way.

OTHER TARGETS Although this review has emphasized spectra of the nucleic acid type, spectra that point to other molecules are common in the literature. For the sake of brevity, I will cite a few without comment. Ultraviolet AS exist for membrane dam- age (62,631, alanine uptake (64), growth delay (65) and loss of saxitoxin binding sites (66).

SOME EFFECTS NOT INCLUDED Due to space limitations, I did not review studies that con- cerned photoreactivation, indirect photoreactivation, photo- protection, photomorphogenesis, production of specific dam- age to nucleic acids, many functions of higher plants, reac- tions of human skin other than those mentioned above, al- most all of the recent literature that concerns field studies or polychromatic exposure and only mentioned in passing some effects in the UVA. Other omissions are too numerous to mention.

Considerations of the limits of interpretation, methods of analysis and the appropriate conditions for the application of action spectroscopy are available (67-73).

REFERENCES 1. Daubeny. C. (1836) On the action of light upon plants and

plants upon the atmosphere. Phil. Trans. R. Soc. London, pp. 149-175.

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Photochemistry and Photobiology, 1997, 65s 127s

2. Senebier, J. ( 1785) Physikalisch-chemische abhandlungen uber den einfluss des sonnenlichtes auf alle drei reiche der natur und auf das pflanzen-reich in sonderheit. Pub. Jacobeer, Leipzig.

3. Englemann, T. W. (1882) Uber sauerstoffausscheidung von pfllanzensellen im microspectrum. Bot. Zeit. 40, 419426.

4. Draper, J. W. (1884) Note on decomposition of carbonic acid ' by the leaves of plants under the influence of yellow light. Phil.

Mag. Ser. 3 25, 169-173. 5. Hausser, K. W. and W. Vahle (1931) Die abhangigkeit des lich-

terythems und der pigmentbildung von der schwingungszahl (wellenlange der erregenden strahlung). Strahlentherapie 13,

6. Berger, D.. F. Urbach and R. E. Davies (1968) The action spec- trum of erythema induced by ultraviolet radiation, preliminary report. XI11 Con-gressus internationalis dermatologiae, Munch- en 1967, pp. 1 1 12-1 117. Springer-Verlag, Berlin, New York.

7. Coblentz, W. W., R. Stair and J. M. Hogue (1931) The spectral erythemic reaction of the human skin to ultraviolet radiation. Proc. Natl. Acad. Sci. USA 17, 401-405.

8. Luckiesh, M. (1930) Artificial sunlight. Trans. Illum. Eng. Soc. 26, 77.

9. Hausser, K. W. and W. Vahle (1927) Sonnenbrand und Son- nenbraunung. Wiss. Veroff Siemens Konzern 6, 101-120.

10. Hausser, K. W. and W. Vahle (1969) Sunburn and suntanning. In The Biologic Effects of Ultraviolet Radiation. (Edited by F. Urbach), pp. 3-21. Pergamon Press, Oxford, New York. [Trans- lated by F. Urbach]

1 1 . Blum, H. F. (1943) Wavelength dependence of tumor induction by ultraviolet radiation. J. Natl. Cancer Inst. 3, 433-537.

12. Forbes, P. D., R. E. Davies and F. Urbach (1978) Experimental ultraviolet photocarcinogenesis-wavelength interactions and time-dose relationships. Natl. Cancer Inst. Monogr. 50, 3 1-38.

13. De Gruijl, F. R. and J. C. Van der Leun (1994) Estimate of the wavelength dependency of ultraviolet carcinogenesis in humans and its relevance to the risk assessment of a stratospheric ozone depletion. Health Phys. 67, 320-325.

14. Setlow, R. B. (1974) The wavelengths in sunlight effective in producing skin cancer: a theoretical analysis. Proc. Natl. Acad. Sci. USA 71, 3363-3366.

15. Setlow, R. B., E. Grist, K. Thompson, and A. Woodhead (1993) Wavelengths effective in induction of malignant melanoma. Proc. Narl. Acad. Sci. USA 90, 6666-6670.

16. Harris, F. I. and H. S. Hoyt (1917) The possible origin of the toxicity of ultraviolet light. Science 46, 3 18-320.

17. Gates, F. L. (1930) A study of the bactericidal action of ultra- violet light. 111. The absorption of ultraviolet light by bacteria. J. Gen. Physiol. 14, 31-42.

18. Gates, F. L. (1928) On nuclear derivatives and the lethal action of ultraviolet light. Science 68, 479480.

19. Gates, F. L. (1929) A study of the bactericidal action of ultra- violet light. 11. The effect of various environmental conditions. J. Gen. Physiol. 13, 249-260.

20. Gates, F. L. (1929) A study of the bactericidal action of ultra- violet light. I. The reaction to monochromatic radiations. J. Gen. Physiol. 13, 23 1-248.

21. Gates, F. L. ( 1934) Results of irradiating Staphylococcus aureus bacteriophage with monochromatic ultraviolet light. J. Exp. Med. 60, 179-188.

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