A 'misplaced chapter' in the history of photosynthesis research; the second publication (1796) on plant processes by Dr Jan Ingen-Housz,MD, discoverer of photosynthesis

In 1779, the Dutch physician Jan Ingen-Housz (1730–1799) obtained a leave-of-absence from his post as Court Physician to Empress Maria Theresa of Austria in order to do research (in England) on plants during the summer months. He performed more than 500 experiments, and described the results in his exceptional book Experiments Upon Vegetables (1779). In addition to proving the requirement for light in photosynthesis, Ingen-Housz established that leaves were the primary sites of the photosynthetic process. Later, Ingen- Housz published research papers on various subjects but aside from his 1779 book, he published only one more communication on photosynthesis and plant physiology. This was entitled 'An Essay on the Food of Plants and the Renovation of Soils'. The essay was published in 1796 as an appendix to an obscure British government report, which is rare and virtually unknown. The present paper describes the 1796 essay, which is particularly interesting in that it shows how Ingen-Housz's concepts were modified by new interpretations of chemical phenomena described in Lavoisier's great and revolutionary book Traité Élémentaire de Chimie (1789). Ingen-Housz not only discovered photosynthesis, but plant respiration as well, and the 1796 essay is testimony to his remarkable insights.     

An integration of photosynthetic traits and mechanisms that can increase crop photosynthesis and grain production

The hypothesis we propose is that during photosynthesis the balance between potentially detrimental and beneficial photochemically induced events can be tipped beneficially toward increased photosynthesis and toward increased crop yield. To test this hypothesis a procedure has been devised with the rice plant, Oryza sativa, that has resulted in increasing both canopy photosynthesis and rice grain yield. Two elite rice varieties selected independently in the contrasting environments of either South China or Texas, each with distinct photosynthetic traits, were crossed to produce a hybrid with an increased canopy photosynthesis and grain yield that is regularly 20 to 22% higher than the mid-yields of the parents. The photosynthetic and mechanisms which may contribute to these beneficial results in the hybrid rice are: a reduction of the midday depression of photosynthesis; a rapid development of the canopy for photosynthetic light interception and an increased canopy photosynthesis; increased amounts of carotenoids for the xanthophyll cycle; an increased protection against free radicals induced by paraquat treatment; a 6 to 12 day shorter plant reproductive life cycle; and a 8 to 10 day increase in the longevity of the flag leaf over the parents. While the hybrid rice has successfully integrated these and likely other unknown characteristics to increase both crop photosynthesis and grain yield, we propose that understanding the underlying beneficial photosynthetic mechanisms supporting these crop plant traits is worthy of thorough investigation and application in crop production.Dedicated to the memory of Professor D.I. Arnon who enriched and challenged the study of photosynthesis through a series of discoveries over 4 decades and via his force of personality.     

Martin Gibbs (1922–2006): Pioneer of 14C research, sugar metabolism & photosynthesis; vigilant Editor-in-Chief of Plant Physiology; sage Educator; and humanistic Mentor

The very personal touch of Professor Martin Gibbs as a worldwide advocate for photosynthesis and plant physiology was lost with his death in July 2006. Widely known for his engaging humorous personality and his humanitarian lifestyle, Martin Gibbs excelled as a strong international science diplomat; like a personal science family patriarch encouraging science and plant scientists around the world. Immediately after World War II he was a pioneer at the Brookhaven National Laboratory in the use of ¹⁴C to elucidate carbon flow in metabolism and particularly carbon pathways in photosynthesis. His leadership on carbon metabolism and photosynthesis extended for four decades of working in collaboration with a host of students and colleagues. In 1962, he was selected as the Editor-in-Chief of Plant Physiology. That appointment initiated 3 decades of strong directional influences by Gibbs on plant research and photosynthesis. Plant Physiology became and remains a premier source of new knowledge about the vital and primary roles of plants in earth’s environmental history and the energetics of our green-blue planet. His leadership and charismatic humanitarian character became the quintessence of excellence worldwide. Martin Gibbs was in every sense the personification of a model mentor not only for scientists but also shown in devotion to family. Here we pay tribute and honor to an exemplary humanistic mentor, Martin Gibbs.     

The construction of a scientific model: Otto Warburg and the building block strategy

In the years 1919 to 1923, Otto Warburg published four papers that were to revolutionise the field of photosynthesis. In these articles, he introduced a number of new techniques to measure the rate of photosynthesis, put forward a new model of the mechanism and added a completely new perspective to the topic by attempting to establish the process’s efficiency in terms of the light quantum requirement. In this paper I trace the roots of Warburg’s series of contributions to photosynthesis research by exploring three different contexts of inspiration: Warburg’s own research into cell respiration, his father’s work on the quantum yield of photochemical reactions in general and the photosynthesis work carried out by Richard Willstätter and Arthur Stoll. When these influences are considered together, it becomes clear that Warburg implemented a Building Block Strategy in his research: rather than inventing his photosynthesis model from scratch, he availed himself of fragments from other contexts, which he then recombined in a new and innovative way. This way of working is considered to be standard practice in scientific research.     

Photosynthesis and carbohydrate metabolism in tobacco leaves during an incompatible interaction with Phytophthora nicotianae

The metabolic and cellular changes in source leaves of Nicotiana tabacum L. cv SNN during an incompatible interaction with Phytophthora nicotianae van Breda de Haan were investigated and compared with defence reactions. Hypersensitive cell death was preceded by a rapid and highly localized shift to non-assimilatoric metabolism. During the first 6 h post infection (hpi), reactive oxygen species (ROS) accumulated. Callose was deposited at the interface of adjacent mesophyll cells (≥1 hpi), the export of sucrose collapsed and its content in the apoplast increased. Stomata closed and photosynthetic flux was reallocated from CO₂ assimilation in favour of photorespiration. This was accompanied by an increase in respiration, glucose-6-phosphate dehydrogenase (G6PDH) activity, apoplastic invertase and hexose content. Later (>6 hpi) the photosynthetic electron transport chain was interrupted and photosynthesis completely collapsed. This was accompanied by a further increase in apoplastic invertase and carbohydrates, respiration and oxidative pentose phosphate pathway (OPPP) and followed by further burst in ROS release. Hypersensitive cell death did not appear until photosynthesis completely declined. Photosynthesis was visualized by chlorophyll-a fluorescence imaging on a macro- and microscopic scale. Decline in photosynthesis and defence reactions were highly localized processes, which occur in single mesophyll cells. We propose that in photoautotrophic leaves, photosynthesis and assimilatory metabolism must be switched off to initiate respiration and other processes required for defence. An early blockage of intercellular sugar transportation, due to callose deposition, in conjunction with enhanced apoplastic invertase activity could facilitate this metabolic shift.     

Changes in photosynthetic activity of buds and stem tissues of Fagus sylvatica during winter

Summary The photosynthetic activity of leafless twigs and buds of Fagus sylvatica was determined by in vivo chlorophyll fluorescence from November to May. Measurements were made on the day of sampling, and during exposure to warm temperatures until reactivation was attained. Under the same conditions, bud development and growth were forced by exposure of cut twigs to 25/18° C at long-day conditions, and bud swelling and bud burst were monitored. Winter inactivation of photosynthesis results in a reduction of the photochemical efficiency of PS II, as indicated by lowering of F_V/F_M, from January through March. The greatest reduction is in cortical chlorenchyma, the least in folded leaflets and primordia of buds. Restoration of photosynthetic activity, brought about by warming, needed 3–4 weeks in cortex and 1–2 weeks in buds during the coldest period of winter. Frequency distributions based on three types of chlorophyll fluorescence transients, defined by quantitative fluorescence parameters, have proved to be a valuable method for a differentiated expression of the unequal functional activation states of parallel samples. The seasonal course of winter inactivation of photosynthesis did not correspond entirely with the depth of bud dormancy as revealed by the forcing treatments; inactivation of photosynthesis may be more closely synchronized with changes in frost hardiness; possible causes are discussed. We suggest, therefore, that a distinction should be made between inactivation of metabolic processes and depth of dormancy, even though these processes are inherently interrelated.Dedicated to Professor Otto Härtel on the occasion of his 80th birthday     

Charles Stacy French: A tribute

Charles Stacy French, one of the great men of photosynthesis research, died on 13 October 1995. He received his PhD at Harvard University where he associated with William Arnold, Caryl Haskins, later president of the Carnegie Institution of Washington, and Pei-Sung Tang. He did early work on the photosynthesis of photosynthetic bacteria with Robert Emerson and later with Otto Warburg. French worked for three years with James Franck in Chicago. His associates there included Hans Gaffron, Robert Livingston, Warren Butler and Roderick Clayton. After spending three years at the University of Minnesota he became the director of the Department of Plant Biology of the Carnegie Institution of Washington and remained there until he retired in 1973. French's research career at the Carnegie Institution was marked by the development of novel and ingenious pieces of equipment such as the French pressure cell used to prepare chloroplast particles to measure partial reactions of photosynthesis. He developed the first recording fluorescence spectrophotometer and demonstrated efficient energy transfer between certain photosynthetically-active pigments, a spectrophotometer that measured the first derivative of absorbance, as well as a novel analog computer to show that complex absorption curves in living plants are produced by a number of distinct forms of chlorophyll occurring in vivo. French used the Blinks rate-measuring oxygen electrode to measure action spectra of oxygen evolution by photosynthesis automatically. He and Jack Myers did some of the pioneering work on the Emerson effect showing the necessary cooperation of two photosystems in photosynthesis. French used the Carnegie Institution's fellowship program to bring large numbers of scientists from around the world to his laboratory. When Stacy French died in 1995, the field of photosynthesis lost one of its great and early pioneers.This is CIW/DPB publication No. 1314.     

The legacy of Hans Molisch (1856–1937), photosynthesis savant

Hans Molisch (1856–1937) was an exceptionally gifted and productive researcher who had broad interests in plant biology, physiology and biochemistry. In addition, he pioneered in isolating a number of species of purple photosynthetic bacteria in pure culture (including Rhodobacter capsulatus), which facilitated his discovery of basic aspects of bacterial photosynthesis. Molisch demonstrated conclusively that molecular oxygen is not produced by photosynthetic bacteria, and discovered the photoheterotrophic growth mode. The range of Molisch's research accomplishments was impressive, and he emerges as a major figure in the history of photosynthesis research. This essay reviews the numerous research contributions made by Molisch, particularly in regard to advancing knowledge of the several forms of photosynthetic metabolism. An English translation of his 1914 paper on the photosynthetic creation of visual images on leaves is included as an Appendix.     

Govindjee,an institution,at his 80th (really 81st) birthday in Třeboň in October, 2013: a pictorial essay

Govindjee (one name only), who himself is an institution, has been recognized and honored by many in the past for he is a true ambassador of “Photosynthesis Research” to the World. He has been called “Mr. Photosynthesis”, and compared to the Great Wall of China. To us in T?eboň, he has been a great research collaborator in our understanding of chlorophyll a fluorescence in algae and in cyanobacteria, and more than that a friend of the Czech “Photosynthesis” group, from the time of Ivan ?etlík (1928–2009) and of Zdeněk ?esták (1932–2008). Govindjee’s 80th (really 81st) birthday was celebrated by the Institute of Microbiology, Laboratory of Photosynthesis, by toasting him with an appropriate drink of a suspension of green algae grown at the institute itself. After my presentation, on October 23, 2013, of Govindjee’s contributions to photosynthesis, and his intimate association with the photosynthetikers (in Jack Myers’s words) of the Czech Republic, Govindjee gave us his story of how he began research in photosynthesis in the late 1950s. This was followed by a talk on October 25 by him on “Photosynthesis: Stories of the Past.” Everyone enjoyed his animated talk—it was full of life and enjoyment. Here, I present a brief pictorial essay on Govindjee at his 80th (really 81st) birthday in T?eboň during October 23–25, 2013.     

Water-use efficiency as a means of modelling net assimilation in boreal forests

Although the processes governing photosynthesis are well understood, scaling from shoot to canopy in coniferous forests is complex. Development of different sap-flow techniques has made it possible to measure transpiration of whole trees and thereby also of whole canopies. There is a strong link between photosynthesis and transpiration, for which reason it would be interesting to test whether measurements of canopy transpiration could also be used to estimate canopy photosynthesis. As a first step towards this, water-use efficiency (WUE) was studied at branch and canopy scales on the basis of branch gas-exchange measurements, with half-hourly and daily temporal resolution. Half-hourly and daily WUE at both branch and canopy scales showed a strong dependency on vapour-pressure deficit ('e). Branch photosynthesis modelled from branch transpiration and 'e mimicked well measured branch photosynthesis. Also, modelled photosynthesis, scaled to canopy and compared to net forest CO₂ exchange measured by the eddy-covariance technique, occasionally showed good agreement. In spite of these seemingly promising results, there was a difference in the response to 'e between branches and between years, which needs to be better understood.     

Light-driven quinone reduction in heliobacterial membranes

Vyacheslav Vasilevich (V.V.) Klimov (or Slava, as most of us called him) was born on January 12, 1945 and passed away on May 9, 2017. He began his scientific career at the Bach Institute of Biochemistry of the USSR Academy of Sciences (Akademy Nauk (AN) SSSR), Moscow, Russia, and then, he was associated with the Institute of Photosynthesis, Pushchino, Moscow Region, for about 50 years. He worked in the field of biochemistry and biophysics of photosynthesis. He is known for his studies on the molecular organization of photosystem II (PSII). He was an eminent scientist in the field of photobiology, a well-respected professor, and, above all, an outstanding researcher. Further, he was one of the founding members of the Institute of Photosynthesis in Pushchino, Russia. To most, Slava Klimov was a great human being. He was one of the pioneers of research on the understanding of the mechanism of light energy conversion and of water oxidation in photosynthesis. Slava had many collaborations all over the world, and he is (and will be) very much missed by the scientific community and friends in Russia as well as around the World. We present here a brief biography and some comments on his research in photosynthesis. We remember him as a friendly and enthusiastic person who had an unflagging curiosity and energy to conduct outstanding research in many aspects of photosynthesis, especially that related to PSII.     

Seasonal and diurnal changes in photosynthetic limitation of young sweet orange trees

This study tests the hypothesis that potted sweet orange plants show a significant variation in photosynthesis over seasonal and diurnal cycles, even in well-hydrated conditions. This hypothesis was tested by measuring diurnal variations in leaf gas exchange, chlorophyll fluorescence, leaf water potential, and the responses of CO₂ assimilation to increasing air CO₂ concentrations in 1-year-old ‘Valência’ sweet orange scions grafted onto ‘Cleopatra’ mandarin rootstocks during the winter and summer seasons in a subtropical climate. In addition, diurnal leaf gas exchange was evaluated under controlled conditions, with constant environmental conditions during both winter and summer. In relation to our hypothesis, a greater rate of photosynthesis is found during the summer compared to the winter. Reduced photosynthesis during winter was induced by cool night conditions, as the diurnal fluctuation of environmental conditions was not limiting. Low air and soil temperatures caused decreases in the stomatal conductance and in the rates of the biochemical reactions underlying photosynthesis (ribulose-1,5-bisphosphate (RuBP) carboxylation and RuBP regeneration) during the winter compared to the values obtained for those markers in the summer. Citrus photosynthesis during the summer was not impaired by biochemical or photochemical reactions, as CO₂ assimilation was only limited by stomatal conductance due to high leaf-to-air vapor pressure difference (VPD) during the afternoon. During the winter, the reduction in photosynthesis during the afternoon was caused by decreases in RuBP regeneration and stomatal conductance, which are both precipitated by low night temperature.     

In memory of Achim Trebst (1929–2017): a pioneer of photosynthesis research

The life and work of Achim Trebst (1929–2017) was dedicated to photosynthesis, involving a wide span of seminal contributions which cumulated in more than five decades of active research: Major topics include the separation of light and dark phases in photosynthesis, the elucidation of photosynthesis by the use of inhibitors, the identification of the three-dimensional structure of photosystem II and its degradation, and an explanation of singlet oxygen formation. For this tribute, which has been initiated by Govindjee, twenty-two personal tributes by former coworkers, scientific friends, and his family have been compiled and combined with an introduction tracing the different stages of Achim Trebst’s scientific life.     

On the requirement of minimum number of four versus eight quanta of light for the evolution of one molecule of oxygen in photosynthesis: A historical note

A bitter controversy had existed as to the minimum number of quanta required for the evolution of one molecule of oxygen in photosynthesis: Otto Warburg had insisted since 1923 that this value was 3–4, whereas Robert Emerson and others continued to obtain a value of 8–12 since the 1940s. It is shown in this letter that the 1931 Nobel-laureate of Physiology & Medicine Otto Warburg published, in his last and final paper, just before his death in 1970, a measured minimum quantum requirement of oxygen evolution of 12 at the lowest intensities of light he used. Although using his theory on photolyte, Warburg calculated a value of 3–4 for the quantum requirement, this is the first confirmation by Warburg of the higher measured quantum requirement. However, it has remained unknown to most investigators. It is expected that this information will be of general interest not only to those interested in the history and research on photosynthesis, but to the entire sci entific community, especially the writers of text books in biology, biochemistry and biophysics.     

Optimal Control of Gas Exchange during Drought: Empirical Evidence

The optimal regulation model by Mäkelä, Berningerand Hari (Annals of Botany 77: 461–467, 1996) was appliedto data for photosynthesis and transpiration of Scots pine duringa 22-d drought period. There was a clear decrease in photosynthesisand transpiration during that period. The agreement betweenmodel and photosynthesis data was good. The residuals of photosynthesiswere not systematic with respect to temperature, irradianceor water vapour deficit. However, the model initially overestimatedtranspiration by 50%, although there was a clear linear relationshipbetween measured and estimated values. The results suggest thatthere was no decrease in photosynthetic capacity during theperiod, but a decrease in stomatal conductance was responsiblefor the changes in photosynthesis and transpiration. The observationsare similar to results in the literature. Transpiration; photosynthesis; stomatal conductance; drought; Pinus sylvestris     

High-temperature inhibition of photosynthesis is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survival of dipterocarp seedlings in the understorey of south‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. Field measurements demonstrated that Shorea leprosula seedlings in a rain forest understorey received a high proportion of daily photon flux density at temperatures supra‐optimal for photosynthesis (72% at ≥30 °C, 14% at ≥35 °C). To investigate the effect of high temperatures on photosynthesis during sunflecks, gas exchange and chlorophyll fluorescence measurements were made on seedlings grown in controlled environment conditions either, under uniform, saturating irradiance (approximately 539 µmol m^?2 s^?1) or, shade/fleck sequences (approximately 30 µmol m^?2 s^?1/approximately 525 µmol m^?2 s^?1) at two temperatures, 28 or 38 °C. The rate of light‐saturated photosynthesis, under uniform irradiance, was inhibited by 40% at 38 °C compared with 28 °C. However, during the shade/fleck sequence, photosynthesis was inhibited by 59% at 38 °C compared with 28 °C. The greater inhibition of photosynthesis during the shade/fleck sequence, when compared with uniform irradiance, was driven by the lower efficiency of dynamic photosynthesis combined with lower steady‐state rates of photosynthesis. These results suggest that, contrary to current dogma, sunfleck activity may not always result in significant carbon gain. This has important consequences for seedling regeneration processes in tropical forests as well as for leaves in other canopy positions where sunflecks make an important contribution to total photon flux density.