Last days in the old radiation laboratory (ORL), Berkeley,California, 1954

Govindjee, the founding editor of the Historical Corner of Photosynthesis Research, invited me 3 years ago to tell the story of why I left Melvin Calvin’s laboratory in the mid 1950s long before the 1961 Nobel Prize in Chemistry was awarded to Calvin for the path of carbon in photosynthesis. I have already written my scientific perspective on this topic (see Benson (Photosynth Res 73:29–49, 2002); also see Bassham (Photosynth Res 76:35–52, 2003) as he was also a major player in this research). Here, I present my recollections of my last days in the old radiation laboratory (ORL) at Berkeley, California. References have been added by Govindjee for the benefit of the readers.     

Martin Gibbs and the peaceful uses of nuclear radiation, <Superscript>14</Superscript>C

This abstract is a prologue to this paper. Prior to his health failing, Martin Gibbs began writing remembrances of his education and beginning a science career, particularly on the peaceful uses of nuclear radiation, at the U.S. Brookhaven National Laboratory (BNL), Camp Upton, NY. Two years before his death Martin provided one of us (Govindjee) a draft text narrating his science beginnings in anticipation of publication in Photosynthesis Research. Govindjee edited his draft and returned it to him. Later, when it became difficult for him to complete it, he phoned Govindjee and expressed the desire that Govindjee publish this story, provided he kept it close to his original. Certain parts of Martin’s narrations have appeared without references (Gibbs 1999). The Gibbs family made a similar request since the narrations contained numerous early personal accounts. Clanton Black recently presented an elegant tribute on Martin Gibbs and his entire science career (Black 2008). Clanton was given the draft, which he and Govindjee then agreed to finish. This chronicle is their effort to place Gibbs’s narrations about his education and his maturation scientifically, in context with the beginnings of biological chemistry work with carbon-14 at the BNL (see Gibbs 1999). Further, these events are placed in context with those times of newly discovered radioisotopes which became available as part of the intensive nuclear research of World War II (WW II). Carbon-14, discovered during WW II nuclear research in 1940, was extremely useful and quickly led to the rapid discovery of new carbon metabolism pathways and biochemical cycles, e.g., photosynthetic carbon assimilation, within a decade after WW II.

A list of personal perspectives with selected quotations,along with lists of tributes,historical notes,Nobel and Kettering awards related to photosynthesis

The history of photosynthesis research can be found in original papers and books. However, a special history is available from the prefatory chapters and the personal perspectives of various researchers who published them in several journals over the last 40 years. We have compiled a list of such perspectives published since 1964. Selection is not easy, especially of authors who were not directly engaged in photosynthesis research; some are included for their special insights related to central issues in the study of photosynthesis. Our journal, Photosynthesis Research, contains other valuable historic data in the occasional tributes, obituaries and historical notes, that have been published. Lists of these items are included. This article ends by listing the Nobel prizes related to photosynthesis and the Kettering Awards for Excellence in Photosynthesis Research. Wherever possible, a web page address is provided. The web page addresses have been taken from the article `Photosynthesis and the Web: 2001' by Larry Orr and Govindjee, available at http://www.life.uiuc.edu/govindjee/photoweb and at http://photoscience.la.asu.edu/photosyn/ photoweb/default.html. ‘When I find a bit of leisure I trifle with my papers. This is one of the lesser frailities. – Horace, Satires I, IV. This revised version was published online in June 2006 with corrections to the Cover Date.     

List of biography and history published mostly in <Emphasis Type="Italic">Photosynthesis Research</Emphasis>, 1988–2008

As an outgoing Editor of the Historical Corner of Photosynthesis Research, I present here the following list of papers of historical interest for the benefit of all. The first paper I published was: Govindjee (1988) The Discovery of Chlorophyll–protein Complex by Emil L. Smith during 1937–1941. Photosynth Res 16:285–289. In order to bring to the readers this List of references on the historical papers published in this journal (and some even elsewhere), I have organized these papers under the following headings (some are arbitrarily assigned to a particular section since they may fit in more than one section): (I) biographies (that include obituaries and tributes, arranged alphabetically, with dates of birth and death); (II) recognitions of scientists (arranged alphabetically) by others; (III) personal perspectives (arranged alphabetically); (IV) historical papers (first chronologically, by the year of publication, and then alphabetically by the names of the editors); (V) special issues of Photosynthesis Research (chronologically by the year of publication and then alphabetically by the names of editors); and lastly (VI) Conferences (available reports in Photosynthesis Research). I will appreciate readers to send me (by e-mail: gov@illinois.edu) corrections, if any, and additional references from other journals. “The firefly seems afire, the sky looks flat; Yet sky and fly are neither this nor that”     

Origin and evolution of C4 photosynthesis in the tribe Salsoleae (Chenopodiaceae) based on anatomical and biochemical types in leaves and cotyledons

 The Chenopodiaceae genus Salsola contains a large number of species with C₄ photosynthesis. Along with derivative genera they have a prominent position among the desert vegetation of Asia and Africa. About 130 species from Asia and Africa were investigated to determine the occurrence of C₃ versus C₄ syndrome in leaves and cotyledons, and to study specific anatomical and biochemical features of photosynthesis in both photosynthetic organs. The species studied belong to all six previously identified sections of the tribe Salsoleae based on morphological characters. Types of photosynthesis were identified using carbon ¹³C/¹²C isotope fractionation. The representatives of all systematic groups were investigated for mesophyll anatomy and biochemical subtypes by determination of enzyme activity (RUBPC, PEPC, NAD- and NADP-ME and AAT) and primary photosynthetic products. Two photosynthetic types (C₃ and C₄) and two biochemical subtypes (NAD- and NADP-ME) were identified in both leaves and cotyledons. Both Kranz and non-Kranz type anatomy were found in leaves and cotyledons, but cotyledons had more diversity in anatomical structure. Strong relationships between anatomical types and biochemical subtypes in leaves and cotyledons were shown. We found convincing evidence for a similar pattern of structural and biochemical features of photosynthesis in leaves and cotyledons within systematic groups, and evaluated their relevance at the evolutionary level. We identified six groups in tribe Salsoleae with respect to photosynthetic types and mesophyll structure in leaves and cotyledons. Two separate lineages of biochemical and anatomical evolution within Salsoleae were demonstrated based on studies of leaves and cotyledons. The sections Caroxylon, Malpighipila, Cardiandra and Belanthera have no C₃ species and only the NAD-ME C₄ subtype has been found in leaves. We suggest the C₄ species in the NADP-ME lineage evolved in Coccosalsola and Salsola sections, and originated in the subsection Arbuscula. Coccosalsola contains many species with C₃ and/or C₃-C₄ intermediate photosynthesis. Within these main evolutionary lineages, species of different taxonomic groups (sections and subsections) had differences in anatomical or/and biochemical features in leaves and cotyledons. We conclude that structural and biochemical changes in the photosynthetic apparatus in species of the tribe Salsoleae were a key factor in their evolution and broad distribution in extreme desert environments. Received January 25, 2001 Accepted July 17, 2001     

Phylogeny, evolution and biogeography of the Quadrifoliovariinae Yamaguti, 1965 (Digenea: Lecithasteridae)

The Quadrifoliovariinae is revised and three new species of Quadrifoliovarium Yamaguit, 1965 from acanthurid fishes of the genus Naso from waters of the Indo-Pacific are described: Q, maceria n. sp. from N.␣tonganus, N. annulatus, N. fageni and N. brevirostris; Q. simplex n. sp. from N. tonganus and N.␣annulatus; and Q. quattuordecim n. sp. from N. tonganus. Amendments are made to the characterisation of the Quadrifoliovariinae, Quadrifoliovarium, Bilacinia Manter, 1969 and Unilacinia Manter, 1969 in light of observations on type and new material. A molecular phylogeny based on ITS2 and 28S regions of the ribosomal DNA is proposed. The phylogeny suggests that U. asymmetrica is the most basal taxon and Q.␣simplex n. sp. and Q. quattuordecim n. sp. the most derived. Evolution of morphological traits within the Quadrifoliovariinae are discussed in light of the molecular phylogeny. Molecular sequences of the ITS2 rDNA were identical between specimens of Q. pritchardae collected off Exmouth (Indian Ocean), Heron Island and Lizard Island (Western Pacific) and Moorea (far Eastern Indo-Pacific), indicating a broad Indo-Pacific distribution. All members of the subfamily are recorded only from the acanthurid genus Naso, with the exception of B. lobatum (Yamaguti, 1970), which has been recorded from a pomacanthid. The restricted host range of the group is discussed in the light of the phylogeny of the host genus Naso.     

Photosynthetic activity of epilithic algal communities in sections of the Torna stream (Hungary) with natural and modified riparian shading

The photosynthetic activity of epilithon grown in the Torna-stream (Hungary) was studied. Samples were collected from natural sections with closed canopy and natural stream bed and from modified sections with trapezoid river bed and shores lacking riparian vegetation. We assumed that human modifications would have a significant impact on epilithic photosynthesis, and therefore expected to observe corresponding changes in photosynthesis–irradiance (P–I) parametric. The laboratory P–I measurements were carried out monthly between April 2008 and September 2009 with epilithon after 3 weeks of in situ colonization. The maximum rate of the photosynthesis (P _max) correlated positively with the chlorophyll a content of the samples. Natural sites had lower biomass and P _max values than modified sites, and the biomass differed substantially in spring and summer, the P _max differed significantly only in spring. Natural sections had higher biomass-specific photosynthesis values than modified sections in all seasons, but the difference was significant only in summer–autumn: in summer the samples from the natural sections had significantly higher photosynthetic efficiency. In spring and summer, the photoadaptation parameters (I _k ) of communities grown in modified sections were substantially higher than in natural sections. Light availability appeared to be the major factor regulating the seasonal photosynthetic activity of epilithic communities.     

Twenty years of biophysics of photosynthesis in Padova, Italy (1984–2005): A tale of two brothers

This paper tells the history of two brothers, almost a generation apart in age, who met again, after having followed different academic paths, to introduce biophysical research in photosynthesis at the University of Padova. The development of two research groups, one in the Chemistry Department, the other in the Biology Department led to a comprehensive interdisciplinary group across academic barriers. The group of Giovanni Giacometti developed in Physical Chemistry, during the years before his retirement, with some roots which can be traced to the famous Linus Pauling school of the mid 1950s, and made possible, by the work of many students (especially Donatella Carbonera and Marilena Di Valentin) and of an older associate (Giancarlo Agostini). The group participated quite actively with a number of European and American laboratories in the application of physical techniques, especially Electron Spin Resonance (EPR) associated with Optical Spectroscopy (Optically Detected Magnetic Resonance; ODMR), and contributed to the development of the understanding of the structure-function relationships in photosynthetic membrane complexes, stimulated by the determination of the X-ray structure of the purple photosynthetic reaction center in the mid 1980s ( J. Deisenhofer, H. Michel, R. Huber and others). The younger brother of Giovanni, Giorgio Mario Giacometti, came to Padova after obtaining biochemical knowledge from the Rossi-Fanelli school in Rome, where Jeffries Wyman, Eraldo Antonini and Maurizio Brunori were the world masters of hemoglobin research. In Padova, together with a group of young scientists (at first Roberto Bassi and Roberto Barbato, now leaders of their own groups in Verona and in Alessandria respectively, followed soon by brilliant coworkers such as Fernanda Rigoni, Elisabetta Bergantino and more recently Ildikò Szabò and Paola Costantini), Giorgio approached more biochemical themes of oxygenic photosynthesis, such as purification and characterization of antenna chlorophyll-protein complexes, Photosystem II (PS II) particles and subunits, having always in mind structural and molecular problems at the level of the largest integrated particles, which are more difficult to investigate in detail by the spectroscopic techniques.This paper is dedicated to the late Arnold J. Hoff; it may be considered Part 2 of ‘Photosynthesis Research in Italy’, and Giorgio Forti et al.’s (2006) paper as Part 1.     

Dependence of the aboveground CO2 exchange rate on tree size in field-grown hinoki cypress (Chamaecyparis obtusa)

The CO₂ exchange of the aboveground parts for five different-sized 17-year-old (as of 1991) hinoki cypress (Chamaecyparis obtusa) trees growing in the field was non-destructively measured over one year, using an open CO₂ exchange system. The CO₂ exchange of individual trees decreased with decreasing tree sizes, such as aboveground phytomass, leaf mass and leaf area. However, the CO₂ exchange abruptly decreased near the smallest-suppressed sample tree. The size dependence was well described by a generalized power function. The annual gross photosynthesis of individual trees was proportional to the square root of leaf mass or leaf area. The dependence of CO₂ exchange on annual phytomass increment was described by a simple power function with an exponent value less than unity, suggesting that CO₂ exchange per unit of phytomass increment was lower in larger-sized trees than in smaller-sized trees. The mean photosynthetic activity of a tree, i.e., gross photosynthesis per unit of leaf area, slightly increased to its highest value with decreasing leaf area and then decreased abruptly near the smallest sample tree. The maximum value of mean photosynthetic activity was estimated to be 2.85 kg CO₂ m⁻² year⁻¹ for a leaf area of 1.56 m² tree⁻¹. The ratio of mean photosynthetic activity to the maximum photosynthetic activity was the highest in an intermediate tree and decreased gradually toward larger-sized trees by ca. 60% and also decreased toward the smallest suppressed tree by ca. 35%.     

Simulation of photosynthetic plasticity in response to highly fluctuating light: an empirical model integrating dynamic photosynthetic induction and capacity

An empirical model was developed to simulate photosynthetic responses of leaves to highly fluctuating light, with a special focus on the functional role of photosynthetic induction and capacity. Based on diurnal courses of light as input data, which were recorded at natural plant sites, we applied this model to simulate the corresponding course of net photosynthesis (output data) for leaves of two neotropical tree species. All six model input parameters (leaf-specific) were obtained via measurements of leaf gas exchange. The model was tested for leaves in their natural environments, characterized by frequent light-flecks. We compared measured carbon gains with computed ones, using a standard steady-state and our induction model. Simulation runs with the steady-state model can result in an immense overestimation of the true situation, by 13.4% at open sites [pioneer species Heliocarpus appendiculatus (Turczaninow)] and by 86.5% at low light environments of the understorey [mid to late successional species Billia colombiana (Planchon and Lindley)]. These significant overestimations, particularly in the understorey, are mainly the consequence of neglecting a dynamic photosynthetic induction under fluctuating light conditions. The model presented here resulted in clearly improved predictions; in open and understorey sites the true carbon gain of leaves was computed with a mean error of less than 7%. As most leaves at natural plant sites are exposed to light environments allowing for dynamic rather than steady-state CO₂ assimilation, the significance of such induction models is evident and is discussed in relation to scaling-up from leaf to canopy and to the whole plant indicating a large potential for errors. Received: 3 May 1999 / Accepted: 9 July 1999     

Temperature Sensitivity of Photosynthesis in Lolium perenne Swards: A Comparison of Two Methods for Deriving Photosynthetic Parameters from in vivo Measurements

The seasonal variation in photosynthetic rate of grass swards is partly the result of changes in the environment and partly the result of changes in the photosynthetic capacity of the sward itself. We evaluated two types of photosynthesis equations regarding their capacity to analyse seasonal and short-term temperature effects on photosynthesis of ryegrass (Lolium perenne L.). Intact cores of a field-grown ryegrass sward were taken to the laboratory 10 d after cutting for measurement of photosynthesis under controlled conditions. This was done during two four-week periods, in summer and autumn. Net photosynthetic rate (P _N) of the sward was lower in autumn than in summer. Both a simple negatively exponential photosynthesis irradiance-response curve and the Farquhar equations for photosynthesis were applied to the in vivo canopy measurements. Application of the irradiance-response curve showed that irradiance-saturated gross photosynthetic rate increased linearly with increasing temperature and was higher in summer than in autumn. The initial radiation use efficiency did not differ between the seasons but decreased with the temperature rise. This explains the observation that total canopy photosynthetic rate decreased after short-term temperature increases in both seasons. The parameters in Farquhar equations that represent the temperature sensitivity of the maximum electron transport rate and of the Michaelis-Menten constants for CO₂ and O₂ fixation could not be quantified satisfactorily. Parameterisation of the Farquhar equations was hampered by a lack of robust information on many biochemical parameters, and the use of simple empirical response-functions may be preferable in the case of in vivo canopy measurements on grass swards.     

Effect of low water potential on photosynthesis in intact lichens and their liberated algal components

Earlier experiments (T.D. Brock 1975, Planta124, 13–23) addressed the question whether the fungus of the lichen thallus might enable the algal component to function when moisture stress is such that the algal component would be unable to function under free-living conditions. It was concluded that the liberated phycobiont in ground lichen thalli could not photosynthesize at water potentials as low as those at which the same alga could when it was present within the thallus. However, our experience with lichen photosynthesis has not substantiated this finding. Using instrumentation developed since the mid-1970's to measure photosynthesis and control humidity, we repeated Brock's experiments. When applying “matric” water stress (equilibrium with air of constant relative humidity) we were unable to confirm the earlier results for three lichen species including one of the species,Letharia vulpina, had also been used by Brock. We found no difference between the effects of low water potential on intact lichens and their liberated algal components (ground thallus material and isolated algae) and no indication that the fungal component of the lichen symbiosis protects the phycobiont from the adverse effects of desiccation once equilibrium conditions are reached. The photosynthetic apparatus of the phycobiont alone proved to be highly adapted to water stress as it possesses not only the capability of functioning under extremely low degrees of hydration but also of becoming reactivated solely by water vapor uptake.     

Canopy and leaf gas exchange of <Emphasis Type="Italic">Haloxylon ammodendron</Emphasis> under different soil moisture regimes

In order to reveal the drought resistance and adaptation of the C₄ desert plant Haloxylon ammodendron under artificially controlled soil moisture regimes, representative plants were selected to measure canopy photosynthesis using canopy photosynthetic measurement system. The results showed that appropriate soil moisture significantly enhances the canopy and leaf photosynthetic capacity, and extremely high soil moisture is not conducive to the photosynthesis of H. ammodendron.     

Effects of salinity on photosynthetic characteristics in photomixotrophic cell-suspension cultures from Alternanthera philoxeroides

We studied the effect of NaCl salinity on the development of cellular photosynthesis using a green, photomixotrophic, cell-suspension culture of Alternanthera philoxeroides (Mart.) Griseb. For these cells, increasing the concentration of sucrose in the media produces a rapid drop in net photosynthetic rate, which recovers as sucrose is depleted from the media. This predictable recovery provides a simple system to examine cellular photosynthetic development. Cells, unadapted to high salinity, were transferred to nutrient media with 30 mM sucrose (Control) or nutrient media with 30 mM sucrose and 100 mM NaCl (Salt). A dramatic increase in the dark respiration rate of Control and Salt cells during the first 6 d of the experiment produced net oxygen consumption in the light. The high dark respiration rates during this period were accompanied by a decline in total Chl and the amounts of two photosynthetic proteins, the light harvesting Chl a/b binding protein of photosystem II (LHCP) and the small subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco SSU). The dark respiration rate of Salt cells was greater than that of Control cells on days 4–8. After day 4, dark respiration rates decreased and net photosynthesis increased to stable values in both treatments at day 11 after media sucrose concentration reached a minimum. As dark respiration rates decreased and net photosynthetic rates increased, total Chl and the amounts of LHCP and rubisco SSU increased in both Control and Salt cells. The slower development of photosynthetic capacity in salt cells was correlated with a fresh weight that was 20% lower than that of control cells at the end of the experiment.     

植物光合作用的三维特性研究进展

光合作用是地球上最重要的化学反应。虽然针对植物光合作用已经进行了广泛深入的研究,但从三维层面探讨植物叶片光合功能及其调节作用的工作较少。叶片结构、光合机构组分、叶片内光能吸收和传递均具有明显的三维特性,极大影响叶片内CO₂转运、叶肉细胞的电子传递和碳同化,进而使叶片光合功能及其调控表现出复杂的三维特征。因此,从三维角度分析叶片光合特性有助于理解光合作用机理,也能够为提高植物光合作用效率提供理论支持。     

The Rhodopseudomonas viridis photosynthetic membrane: arrangement in situ

The organization of photosynthetic membranes in the cytoplasm of the photosynthetic bacterium Rh. viridis has been examined by several techniques for electron microscopy. Thin sections of membrane stacks show that the regular lattice of membrane subunits reported in other studies can be observed in thin section. Tilting of sections in the electron microscope shows that the regular lattices of several membranes overlap in a way that suggests they are in register with each other. This observation can be confirmed by freeze-fracture images in which a regular arrangement of membrane lattices can be observed, each perfectly aligned.Analysis of the spacings of membrane pairs shows that the photosynthetic membranes of Rh. viridis are very closely apposed. The mean diameter of two membranes is 160A, and the average space between two such membranes is only 42A. When a recently developed atomic level model of Rh. viridis reaction center is superimposed against these spacings, each reaction center extends from the surface of its respective membrane far enough to make contact with an apposing membrane. The limited free space between membranes and regular alignment of lattices has a number of implications for how this membrane is organized to carry out the process of energy transfer.     

I can't believe my luck

The author gives an account of his life and work in scientific research. At the prompting of Govindjee, this is a quite personal account and, I hope, not too serious. The circumstances surrounding the discovery of C₄ photosynthesis are mentioned together with some aspects of the subsequent development of this field. Since it seems to be expected on such occasions, there is also some reminiscences and some unsolicited advice. I hope there are no gross or libelous inaccuracies.Abbreviations 3-PGA 3-phosphoglycerate - PEP phosphoenolpyruvate - P_i orthophosphate - PP_i pyrophosphate Written at the invitation of Govindjee.     

Photosynthetic carbon sources in somepotamogeton species

The ability of somePotamogeton species to use bicarbonate for photosynthesis was assayed by means of relative net photosynthetic rates in alkaline (bicarbonate) and acid (free CO₂) solutions with an equivalent amount of inorganic carbon. The results showedPotamogeton crispus, P. oxyphyllus, P. maackianus, P. perfoliatus, P. malaianus and the submerged leaves ofP. distinctus to be able to use bicarbonate ions for photosynthesis. To the contrary, the submerged leaves ofP. fryeri, which is an inhabitant of soft and acid waters, and the floating leaves ofP. distinctus were proven to be ‘non-users’ of bicarbonate ions. Furthermore, the relative photosynthetic rates were determined using natural waters with various carbon conditions. The dependence of photosynthetic performance on the free CO₂, pH and alkalinity of the waters is discussed in relation to carbon metabolism. Further it is suggested that the carbon conditions are ecologically significant in relation to the productivity and success of macrophyte species in natural habitats.     

Chlorophyll a fluorescence: can it shed light on fundamental questions in photosynthetic carbon dioxide fixation?

Abstract A close, immediate and precise relationship between chlorophyll a fluorescence and photosynthetic carbon assimilation in vivo is demonstrated. The examples discussed include kinetics displayed during dark to light transitions plus oscillations and transients observed during changes in the gas phase surrounding the leaf. Remaining uncertainties surrounding the relationship between chlorophyll fluorescence and photosynthesis are attributed to the underlying complexity of the regulatory mechanisms involved. Examples are also given that show how multiple simultaneous measurements of different aspects of the photosynthetic process may contribute to the resolution of these uncertainties. The practical relevance of these matters is also discussed, particularly in relation to the limitations of the photosynthetic process and to the use of chlorophyll fluorescence as a diagnostic probe of chemical and genetic manipulation and stress.     

Phosphate sequestration by glycerol and its effects on photosynthetic carbon assimilation by leaves

Glycerol induced a limitation on photosynthetic carbon assimilation by phosphate when supplied to leaves of barley (Hordeum vulgare L.) and spinach (Spinacia oleracea L.). This limitation by phosphate was evidenced by (i) reversibility of the inhibition of photosynthesis by glycerol by feeding orthophosphate (ii) a decrease in light-saturated rates of photosynthesis and saturation at a lower irradiance, (iii) the promotion of oscillations in photosynthetic CO₂ assimilation and in chlorophyll fluorescence, (iv) decreases in the pools of hexose monophosphates and triose phosphates and increases in the ratio of glycerate-3-phosphate to triose phosphate, (v) decreased photochemical quenching of chlorophyll fluorescence, and increased non-photochemical quenching, specifically of the component which relaxed rapidly, indicating that thylakoid energisation had increased. In barley there was a massive accumulation of glycerol-3-phosphate and an increase in the period of the oscillations, but in spinach the accumulation of glycerol-3-phosphate was comparatively slight. The mechanism(s) by which glycerol feeding affects photosynthetic carbon assimilation are discussed in the light of these results.Abbreviations Chl chlorophyll - C _i intercellular concentration of CO₂ - P phosphate - PGA glycerate-3-phosphate - Pi orthophosphate - triose-P sum of glyceraldehyde-3-phosphate and dihydroxyacetone phosphate