Bunsen-Roscoe Reciprocity Law Research Articles

Importance of Duration, Duty-Cycling and Thresholds for the Implementation of Ultraviolet C in Marine Biofouling Control

The introduction of a surface into the marine environment begins a process known as biofouling, which increases the weight and hydrodynamic drag of the fouled structure. This process is detrimental to maritime vessels and costs the industry ∼$150B in fuel and maintenance spending annually. Preventing the settlement of fouling organisms mitigates these issues and limits the spread of non-indigenous species (NIS). This is primarily achieved via antifouling paints. Ultraviolet light is a sterilization method used in water purification, food storage packaging, and within medical fields. Ultraviolet C (UV-C) radiation interacts with DNA to prevent growth, proliferation, and survival of bacteria, and biofilm formation. Recent progress in microelectronics technology has advanced the range of commercially available light-emitting diodes (LEDs) to include the UV wavelengths, and the reduced size and cost has allowed their integration into previously inaccessible locales. This study builds on recent progress in integrating UV-C LEDs into UV-lucent silicone tiles for fouling control. The operational cycle needed to prevent growth of Navicula incerta cells was determined. Constant irradiance at a peak of 5.77 μW/cm2 resulted in a significant reduction in diatoms within 2 h, and a 2 log and 3 log reduction after 48 h and 5 days, respectively. Duty cycling (pulsing) in all variations from 50 to 2.5%, indicated significant reductions in cell densities, and the lowest cycle could effectively reduce biofouling growth and increase the longevity of the LEDs for up to 45.6 years. Irradiance and exposure were altered over a set duration and indicated a restriction in growth between 0.01–0.82 J/cm2 and an increased mortality at irradiances > 2.65 J/cm2, suggesting an effective antifouling threshold between these dosages. The effective dosage for 1 log reduction in fouling was estimated to be 25 J/cm2 but varied according to irradiance delivery method. Effective dosage for a 1 log reduction between experimental methods was variable indicating that UV treatment of N. incerta departed from the Bunsen-Roscoe reciprocity law expectancy. The variation in densities at similar dosages could be explained with further investigation of DNA repair mechanisms. In conclusion, UV-B/C use was effective at all irradiances, including as low as 0.01 J/cm2, and holds considerable promise for marine biofouling control.

[Principles of corneal cross-linking : Presentation based on the development of the various treatment protocols].

Corneal cross-linking (CXL) is used to treat corneal ectatic diseases. The aim is to improve the reduced consolidation of the cornea in order to halt further corneal protrusion and therefore subsequent deterioration of the optical imaging proportions. In this article the principles of corneal cross-linking based on riboflavin and UV light are presented including recent research results. Furthermore, the most important treatment protocols including standard CXL (S-CXL), accelerated CXL (A-CXL), transepithelial CXL (TE-CXL) and the approach of the CXL procedure for thin corneas are explained. The CXL method depends on four major components, the riboflavin solution, oxygen, UV light and the availability of cross-linking sites on the collagen tissue. According to the present state of knowledge, the photochemical process of the CXL method induces covalent bonds between the fibrils and proteoglycans and thus stabilizes the collagen fibers, resulting in corneal consolidation. In addition to the S‑CXL, which has proven its effectiveness and safety in alarge number of studies, there are other treatment protocols that have been developed based on the Bunsen-Roscoe law of reciprocity. The A‑CXL protocol has the advantage of having ashorter irradiation time but it seems to be less effective than the S‑CXL protocol concerning the increase in corneal stiffness. The use of TE-CXL has so far not yet gained acceptance in the clinical practice. The CXL procedures primarily aim to stabilize the cornea. In the future, in addition to stabilization of the cornea, simultaneous improvement of visual acuity will be the main focus.

Ultraviolet-B irradiance and cumulative dose combine to determine performance and survival.

Despite decades of research, the role of elevated solar ultraviolet-B radiation (UVBR; 280-315nm) in shaping amphibian populations remains ambiguous. These difficulties stem partly from a poor understanding of which parameters of UVBR exposure - dose, irradiance, and time interval - determine UVBR exposure health risk, and the potentially erroneous assumption that effects are proportional to the dose of exposure, irrespective of the administered regime (Bunsen-Roscoe Law of Reciprocity; BRL). We tested if the BRL holds with respect to UVBR-induced physiological effects in amphibians by acutely exposing tadpoles of the Australian green tree frog (Litoria caerulea) to a combination of different UVBR irradiances and doses in a fully factorial experiment. The BRL was invalid across all metrics assessed, with UVBR irradiance influencing the effects of a given dose on growth, coloration and burst swimming performance of larvae. We demonstrated some of the first empirical evidence for beneficial physiological effects of UVBR exposure in a larval amphibian, with improvements in growth, burst swimming performance and survival at the highest UVBR doses, contrary to hypotheses. Our findings demonstrate the species-specific nature of amphibian responses to UVBR, and the importance of UVBR irradiance in influencing the long-term physiological effects of a given dose of radiation. This work enhances our understanding of which parameters of complex UVBR exposures determine amphibian health risk.

Investigating photodegradation of antibodies governed by the light dosage

The present study investigated the photodegradation of three different monoclonal antibodies (mAb) by visible light. Several chromatographic techniques, such as size-exclusion and hydrophobic interaction chromatography as well as mass spectrometry were used to measure relative changes of various oxidation related monoclonal antibody species. The results show that visible light is indeed capable of inducing the formation of protein photo-oxidation products, such as acidic, basic, hydrophilic, and several other protein species with altered physicochemical properties. Although, the formation rate of degradants of these three protein species was dependent on the light source’s intensity (I), their yield is clearly correlated to the applied light dosage (ld), which is defined as the product of light intensity I and irradiation time t (light dosage = I·t). Hence, our findings indicate that the degradation of monoclonal antibodies can be described according to the Bunsen-Roscoe reciprocity law. This correlation can be useful to assess the impact of photodegradation of biologics with regards to changes in light intensity and/or duration of light exposure of the protein, e.g. during the manufacturing of biologics.

Accelerated corneal collagen cross-linking should be modified.

Dear editor We have read with interest the recently published comparative study regarding the accelerated versus conventional corneal collagen cross-linking (CXL) in the treatment of mild keratoconus. We would like to underline the importance of the CXL treatment time reduction in this time consuming operation.1 Accelerated CXL became popular due to patient and doctor comfort in comparison to the previously widespread use of Dresden protocol which is successful in terms of safety and efficacy. Based on the Bunsen-Roscoe law of reciprocity many modifications to the time and irradiation settings have been proposed while the total energy dose delivered to the cornea should be maintained at 5.4 J/cm2.2 We already proved, by measuring (with the use of anterior segment optical coherence tomography) the demarcation line depth at 1 month postoperatively after CXL, that the treatment depth was significantly different (significantly shallower in accelerated protocol) between the 10 minute accelerated protocol with ultraviolet-A (UV-A) irradiation intensity of 9 mW/cm2 and the 30 minute standard Dresden protocol with UV-A of 3 mW/cm2.2 Our results suggested that Bunsen-Roscoe law of reciprocity may not directly apply to CXL in living cornea tissue, thus, an increased total energy dose should probably be applied to the keratoconic cornea in order to achieve a treatment effect comparable to the already proven effective standard Dresden CXL protocol. Consequently, we have proceeded with relevant modifications for the UV-A time settings of the currently proposed accelerated CXL protocol (10 minutes) by increasing the UV-A irradiation time to 14 minutes (40% increase) and we achieved a treatment effect comparable with the Dresden protocol (as indicated by the demarcation line depth).3 In the recently published article by Sherif, it is presumed that the accelerated group of patients received CXL treatment according to Bunsen-Roscoe law which should have been 3 minutes of UV-A with irradiation intensity of 30 mW/cm2 corresponding to 5.4 J/cm2 total energy dose delivered to the cornea.1 Interestingly, Sherif mentioned that the accelerated group received UV-A treatment with 30 mW/cm2 for 4 minutes and 20 seconds; thus the accelerated group received about 40% more UV-A irradiation time, which is comparable to our time setting modifications (from 10 to 14 minutes). Moreover it should be clarified whether the author modified the time settings manually or if these modifications were made according to the UV-A device manufacturer’s instructions (Avedro, Inc., Waltham, MA, USA). In addition we would like to enquire whether the author performed measurements (eg, endothelial cell density) concerning the safety of the time and irradiation intensity settings used in this study.4 As we previously published, an indirect and non-contact method to measure the effectiveness of CXL treatment is the demarcation line depth provided with anterior segment optical coherence tomography; our results were the same with the reliable but more invasive (due to contact with the treated cornea) confocal microscopy.5 Thus, measurement of the demarcation line depth at 1 month postoperatively could provide useful information in suggesting modifications to any of the CXL treatment settings.6 It should be of great interest whether the author could provide such measurement results concerning the studied groups of this article. Moreover, we should bring to the attention that in cases of mild keratoconus it should probably need a longer follow-up period to establish reliable results concerning the effectiveness of treatment in arresting keratoconus progression. In conclusion we believe that accelerated CXL treatment settings in agreement to Bunsen-Roscoe law of reciprocity should be revised thoroughly and new modifications have to be carefully made regarding time and irradiation intensity adjustments keeping the safety aspect of CXL treatment as the first priority.

The Biomechanical Effect of Corneal Collagen Cross-Linking (CXL) With Riboflavin and UV-A is Oxygen Dependent

In an attempt to reduce treatment time in corneal collagen cross-linking (CXL) with riboflavin and ultraviolet-A (UV-A), recent protocol modifications include shorter irradiation times at higher fluence, while maintaining constant total applied energy (Bunsen-Roscoe law of reciprocity). While such parameter changes might produce similar biological results within a certain range, the limits of reciprocity are unknown. Limitations in the corneal oxygen diffusion capacity and its potential impact on the efficacy of CXL, raise concerns regarding the efficiency of high-fluence CXL, and also of transepithelial CXL. Porcine corneas were treated with an epithelium-off CXL at a fluence of 9 mW/cm2 under two different atmospheres: one with a regular oxygen content (21%) and another in a helium-supplemented, low-oxygen environment (<0.1%). Untreated corneas served as controls (n = 20 each). Five-millimeter corneal stripes were prepared and biomechanical stiffness was measured using an extensometer. Corneas cross-linked under normal oxygen levels showed a significant increase in biomechanical stability (14.36 MPa ± 2.69 SD), whereas corneas treated similarly, but in a low-oxygen atmosphere showed a Young's modulus similar to untreated controls (11.72 MPa ± 2.77 SD). The biomechanical effect of CXL seems to be oxygen dependent. This dependency will be of particular importance in high-fluence and transepithelial CXL and will most likely require major protocol modifications to maintain the efficiency of the method. The oxygen dependency of CXL shown here raises concerns about the effectiveness of high-fluence and transepithelial CXL. Both methods were introduced to clinical ophthalmology without thorough validation.

The Efficacy of Corneal Cross-Linking Shows a Sudden Decrease with Very High Intensity UV Light and Short Treatment Time

Standard treatment in cases of progressive keratectasia is UV-triggered corneal cross-linking. For irradiances larger than 10 mW/cm(2) and treatment times below 10 minutes, the scientific proof of a biomechanical strengthening effect is insufficient. The authors investigated the biomechanical strengthening of ex vivo corneal tissue treated with irradiances between 3 mW/cm(2) and 90 mW/cm(2) and illumination times from 30 minutes to 1 minute, respectively. A total of 100 porcine eyes received riboflavin + UV treatment (constant irradiation dose of 5.4 J/cm(2)) with different intensities and illumination times and were randomly assigned into 10 groups. A control group (80 eyes) was not irradiated but underwent the same treatment otherwise. Young's modulus at 10% strain was determined for each strip after uniaxial stress-strain measurement. A Kruskal-Wallis test was used for statistical analysis. A statistically significant difference (α = 0.01) was found between the median value of Young's modulus of the treatment groups up to 45 mW/cm(2) (illumination times from 30 minutes to 2 minutes) compared with the control group. There was no statistically significant difference between the treatment groups from 50 mW/cm(2) up to 90 mW/cm(2) (illumination times of less than 2 minutes) and the control group. The ex vivo results of corneal cross-linking performed in porcine corneas show that the Bunsen-Roscoe reciprocity law is only valid for illumination intensities up to 40 to 50 mW/cm(2) and illumination times of more than 2 minutes. Further experiments are necessary to validate these results for in vivo human corneal tissue. Additionally, safety aspects at high intensities must be investigated.

The Bunsen–Roscoe reciprocity law in ultraviolet-B-induced mortality of the two-spotted spider mite Tetranychus urticae

To determine whether the Bunsen–Roscoe reciprocity law (i.e., the extent of photochemical effects is determined by cumulative irradiance) is applicable to ultraviolet-B (UVB) damage in the twospotted spider mite Tetranychus urticae, egg hatchability and survival of individuals were assessed after irradiation with a UVB lamp using various combinations of intensity and time length. A positive linear correlation between probit mortality and cumulative UVB irradiance was detected in eggs, larvae, teleiochrysalis females, and adult females, regardless of UVB intensity (0.19–0.58Wm−2). LD50 values were clearly higher in adult females, followed by teleiochrysalis females, larvae, and eggs. In eggs, reciprocity was obeyed not only at the UVB intensities listed above, but also at very low UVB intensity (0.014–0.023Wm−2). Such reciprocity in the negative effects of UVB radiation was also observed for the developmental rate of juveniles and egg production of adult females. However, the LD50 value of eggs obtained using the UVB lamp (0.58kJm−2) was lower than that elicited by solar UVB radiation in a previous outdoor experiment (about 50kJm−2). These results suggest that a photoreactivation mechanism plays an important role in the survival of this mite under solar radiation.

Similarities and differences between phytochrome-mediated growth inhibition of coleoptiles and seminal roots in rice seedlings

In rice, light is known to inhibit the growth of coleoptiles and seminal roots of seedlings through phytochrome. Here we investigated the light-induced growth inhibition of seminal roots and compared the results with those recently determined for coleoptiles. Although three rice phytochromes, phyA, phyB and phyC functioned in a similar manner in coleoptile and seminal root, the Bunsen-Roscoe law of reciprocity was not observed in the growth inhibition of seminal root. We also found coiling of the seminal root at the root tip which appeared to be associated with the photoinhibition of seminal root growth. This could be a new light-induced phenomenon in certain cultivars of rice.

Phytochrome‐mediated growth inhibition of seminal roots in rice seedlings

In rice (Oryza sativa) seedlings, continuous white-light irradiation inhibited the growth of seminal roots but promoted the growth of crown roots. In this study, we examined the mechanisms of photoinhibition of seminal root growth. Photoinhibition occurred in the absence of nitrogen but increased with increasing nitrogen concentrations. In the presence of nitrogen, photoinhibition was correlated with coiling of the root tips. The seminal roots were most photosensitive 48-72 h after germination during the 7-day period after germination. White-light irradiation for at least 6 h was required for photoinhibition, and the Bunsen-Roscoe law of reciprocity was not observed. Experiments with phytochrome mutants showed that far-red light was perceived exclusively by phyA, red light was perceived by both phyA and phyB, and phyC had little or no role in growth inhibition or coiling of the seminal roots. These results also suggest that other blue-light photoreceptors are involved in growth inhibition of the seminal roots. Fluence-response curve analyses showed that phyA and phyB control very low-fluence response and low-fluence response, respectively, in the seminal roots. This was essentially the same as the growth inhibition previously observed at the late stage of coleoptile development (80 h after germination). The photoperceptive site for the root growth inhibition appeared to be the roots themselves. All three phytochrome species of rice were detected immunochemically in roots.

Effect of Intense Pulsed-Light Exposure on Lipid Peroxides and Thymine Dimers in Human Skin In Vivo

Intense pulsed light (IPL) generates high-intensity short flashes of visible light and has been used for about 10 years to improve dermatological conditions such as telangiectasia, pigmented lesions, and skin aging. Although these systems deliver a moderate dose (10-30 J/cm(2)) of visible light, this dose is delivered during a short pulse (2-5 milliseconds), which implies a very high fluence rate (approximately 4000 W/cm(2)). For this reason, we speculated whether the Bunsen-Roscoe law of reciprocity could still be valid in these conditions. Nine healthy volunteers were exposed to IPL or UV-A or simulated solar UV radiation, and then thymine dimer and lipid peroxide concentrations were determined in skin biopsy specimens of the exposed sites. Only exposure to solar UV radiation (7-J/cm(2) UV-A + 80-mJ/cm(2) UV-B) produced measurable amounts of thymine dimers in DNA from skin biopsy specimens, whereas UV-A radiation (40 J/cm(2)) and IPL (9 J/cm(2)) induced 3-fold and 6-fold increases of cutaneous lipid peroxides, respectively. These preliminary results indicate that IPL, although filtered for wavelengths shorter than 500 nm, can generate oxidative stress, a typical hallmark of UV-A, but does not induce thymine dimers. This emphasizes the need for long-term studies involving IPL before using this technique in a recurrent manner.

Asymmetric, blue light-dependent phosphorylation of a 116-kilodalton plasma membrane protein can be correlated with the first- and second-positive phototropic curvature of oat coleoptiles.

The possible correlation between blue light-dependent phosphorylation of a 116-kD protein and phototropic responses of etiolated oat (Avena sativa L.) seedlings was tested by a micromethod for protein phosphorylation. Quantitation of the basipetal distribution of this protein showed that the in vitro 32p phosphorylation values declined exponentially from tip to node, with more than 50% of the total label being found in the uppermost 5 mm. Nonsaturating preirradiation of the coleoptiles in vivo resulted in partial phosphorylation with endogenous ATP. Subsequent in vitro phosphorylation under saturating irradiation allowed the determination of the degree of in vivo phosphorylation. Unilateral preirradiation resulted in higher in vivo phosphorylation on the irradiated than on the shaded side of the coleoptile. The fluence-response curve for the difference in phosphorylation between both sides of the coleoptile resembles the fluence-response curve for first-positive phototropic curvature, although it is shifted by two orders of magnitude to higher fluences. Possible reasons for this shift are discussed. In the coleoptile base the phosphorylation gradient across the coleoptile becomes larger with increasing time of irradiation at a constant fluence. Thus, phosphorylation of the 116-kD protein, in accordance with second-positive phototropic curvature, does not obey the Bunsen-Roscoe reciprocity law.

Solar radiation has a lethal effect on natural populations of culturable outdoor atmospheric bacteria

Ambient heterogenic atmospheric bacteria were collected by impaction directly onto nutrient agar surfaces on clear sunny and cloudy days and at night. Samples were then exposed to summer noontime solar radiation (SR) for increasing periods of time. Bacterial survival was the least for the organisms collected at nighttime and the greatest for those collected during clear sunny days. This result may be due to the inactivation of SR-sensitive bacterial populations by the ambient SR on clear days, and to a lesser extent on cloudy days, leaving only the relatively resistant populations. This does not occur at night when the SR-sensitive populations have not been reduced. The mixed bacterial species populations in the atmosphere do not appear to follow the Bunsen-Roscoe reciprocity law but integration of pure bacterial components of the mixed population that do follow the law, could explain the findings. The populations collected in this investigation exhibited SR survival functions usable in developing survival and dispersion models of outdoor atmospheric bacteria and microbial pesticides.

Blue-Light Regulation of Specific Transcript Levels in Pisum sativum

The expression of many nuclear genes in plants is light regulated. We have examined the fluence-response, time-course, and reciprocity characteristics of four nuclear, blue-light-regulated transcripts in Pisum sativum L. var Alaska: Cab RNA, pEA207 RNA, pEA215 RNA, and pEA25 RNA. To avoid complications due to blue-light-induced transformation of phytochrome, we have adapted the procedure of measuring blue-light-induced changes in steady-state-RNA levels in seedlings grown in continuous red light. The fluence-response curves for Cab and pEA215 RNA are bell-shaped, with peak accumulations at 10(2) and 10(1) micromoles per square meter, respectively. No response is observed at 10(4) micromoles per square meter. pEA25 RNA has threshold and saturation fluences below 10(-1) micromoles per square meter. pEA207 RNA has a threshold at 10(2) micromoles per square meter and saturates above 10(4) micromoles per square meter. pEA215 and Cab RNA start to increase within 1 hour after the 10(1) micromoles per square meter pulse, and finish accumulation by 5 hours. The decrease in pEA207 RNA in response to 10(4) micromoles per square meter and pEA25 RNA in response to 10(1) or 10(4) micromoles per square meter starts between 3 and 5 hours after the blue-light pulse. The Bunsen-Roscoe Law of Reciprocity is followed in the Cab, pEE215, pEA25, and pEA207 RNA responses to 10(1) and 10(4) micromoles per square meter.

Phototropic fluence-response curves for Pilobolus crystallinus sporangiophore

Fluence-response relationships were examined for positive and negative phototropism induced by blue (450 nm) and ultraviolet-B (UV-B, 280 nm) light, respectively, in the Pilobolus crystallinus sporangiophore. Fluence-response curves for both blue and UV-B light obtained by changing the fluence by varying exposure time only showed the classical first and second positive bending. However, fluence-response curves obtained by varying the fluence rate were bell-shaped irrespective of the length of the exposure time. With increasing exposure time the peak became higher along the ascendant arm and the descendant arm was shifted toward the higher fluence. The Bunsen-Roscoe reciprocity law was valid only when the fluence was less than approx. 400 pmol·m(-2) for both blue and UV-B light. Because the shapes of the fluence-response curves for blue and UV-B light were nearly the same, the photoreceptor systems for both blue and UV-B light are considered to be the same.

Photocontrol of Anthocyanin Synthesis

The spectral sensitivity and the irradiance dependence of anthocyanin synthesis, a "high irradiance response," in cabbage (Brassica oleracea, cv. Red Acre) and tomato (Lycopersicon esculentum, cv. Beefsteak) seedlings exposed to continuous irradiation depend upon the length of the exposure. In cabbage, blue and red are more effective than far red when the irradiations are shorter than 12 hours and less effective than far red when the irradiations are longer than 12 hours. The irradiance dependence is negligible under red and becomes evident under blue and far red red only for exposures longer than 12 hours. Anthocyanin synthesis under intermittent light treatments, of efficiency comparable to that of continuous treatments, obeys the Bunsen-Roscoe reciprocity law and is a function of the dose (irradiance x time), rather than of the irradiance alone. The validity of the reciprocity relationships suggests that only one photoreceptor is responsible for the photocontrol of the response in the blue, red, and far red spectral regions. The characteristics of the response suggest that the photoreceptor is phytochrome, at least in cabbage.

Effect of relative humidity on the inactivation of airborne Serratia marcescens by ultraviolet radiation.

Apparatus was designed and constructed in which a bacterial aerosol of known age, particle size, and relative humidity (RH) could be exposed to ultraviolet (UV) radiation of measured intensity for a given period of time and then be sampled quantitatively. Aerosols of Serratia marcescens were exposed to UV dosages between 96.0 and 0.75 (muw-sec)/cm(2) at humidities ranging from 25 to 90%. A sharp decline in the fraction of organisms killed was found at RH values above 60 to 70%. Above 80% RH, there was evidence for reactivation induced by UV. The plot of "log fraction organisms remaining" versus UV dose was curvilinear, suggesting noncompliance with the monomolecular law of reaction velocity, but the Bunsen-Roscoe law of reciprocity between time and intensity of UV exposure was demonstrated to hold. These results could be accounted for by postulating the presence in the aerosol of two populations of organisms with different sensitivities to UV, each individually obeying the monomolecular law of reaction velocity. The data amplify existing information on the relationship between UV disinfection of airborne organisms and RH. In the middle range of humidities, the sensitivity of the organisms to UV was greater than would be expected from published reports.

Effect of Relative Humidity on the Inactivation of Airborne Serratia marcescens by Ultraviolet Radiation

Apparatus was designed and constructed in which a bacterial aerosol of known age, particle size, and relative humidity (RH) could be exposed to ultraviolet (UV) radiation of measured intensity for a given period of time and then be sampled quantitatively. Aerosols of Serratia marcescens were exposed to UV dosages between 96.0 and 0.75 (muw-sec)/cm(2) at humidities ranging from 25 to 90%. A sharp decline in the fraction of organisms killed was found at RH values above 60 to 70%. Above 80% RH, there was evidence for reactivation induced by UV. The plot of "log fraction organisms remaining" versus UV dose was curvilinear, suggesting noncompliance with the monomolecular law of reaction velocity, but the Bunsen-Roscoe law of reciprocity between time and intensity of UV exposure was demonstrated to hold. These results could be accounted for by postulating the presence in the aerosol of two populations of organisms with different sensitivities to UV, each individually obeying the monomolecular law of reaction velocity. The data amplify existing information on the relationship between UV disinfection of airborne organisms and RH. In the middle range of humidities, the sensitivity of the organisms to UV was greater than would be expected from published reports.

THIRD POSITIVE (C-TYPE) PHOTOTROPISM IN THE AVENA COLEOPTILE1

SUMMARY The third (C-type)1 response results from a tonic and a tropistic effect of the irradiation. The Bunsen-Roscoe reciprocity law applies to the tonic as well as to the tropistic component. A simple dependence on the duration of the stimulus does not exist, but is simulated by the time interval required for effectuation of the tonic component. It is concluded that the C-type curvature is identical with the first positive curvature, but is evoked by larger doses of unilateral light because of a desensibilisation of the coleoptiles by the tonic effect of the long unilateral irradiation itself. This desensibilisation process takes a minimum period of 4 resp. 15 minutes to be effective with red-preirradiated or dark-cultivated coleoptiles, respectively.

Morphogenesis in Schizophyllum commune. II. Effects of monochromatic light.

The photoinduction of fruiting bodies by light of defined wavelengths was studied in the fungus Schizophyllum commune Fr. Several properties of the induction were established. (1) The exposure-response relationship for induced fruiting was determined for light of 448 nm. (2) The Bunsen-Roscoe Law of Reciprocity was found to hold for the photoinduction of fruiting bodies for the interval 36 to 2000 sec with light of 448 nm. (3) Light of wavelengths from 320 nm to 525 nm induced fruiting bodies. Although the photoreceptor is unknown, it may be a flavin rather than a carotenoid, because light in the near ultraviolet (350 nm-400 nm) was inductive. (4) Neither red light (660 nm) nor far-red light (730 nm) induced fruiting bodies or affected the sensitivity of the fungus toward photoinduction by blue light (448 nm).