I offer an overview of the contributions made by female scientists to the area of biological photoreceptors (both energy converters and signal-sensing photoreceptors, also called photosensors). In the last few years, there has been a very welcome increase in the participation of women in all areas of research, also in this area. This increase in women’s participation is most certainly due to better children care facilities in many countries and to a better general understanding of the possibilities open to the society by involving men and women equally in all activities including child care.

Therefore, identifying women who were pioneers in this field in the past seems easier than to keep track of all the very productive women who are currently making fundamental inputs. Those presented here were born before 1955 and they are listed following the birthdate. Many women have made extraordinary discoveries. Many others have contributed essential pieces of information through hard and focused work. Some of them are widely remembered, others have been nearly forgotten. Most probably, and due to ignorance, I am leaving out important (she) colleagues. This paper should be considered as a work in progress.

This is not a biographical study of each of the scientists, but I have tried to depict in each case important circumstances that were a turning point for their success.

In most of the biographies of these women, it is pointed out that they supported and encouraged younger female students and scholars and served as path-marking role models. Women scientists tell us in their autobiographical writings about the difficulties they encountered to be accepted as a science student or as a scientist. In the 60s or 70s, some institutions did not enroll women in their graduate courses “because women do not get a Nobel Prize” (see Elaine Tobin, below), which, incidentally, was true!

Birgit Vennesland (Fig. 1). She was born in Norway in 1913. She was a remarkable biochemist who worked in several areas. While studying enzymatic reactions as a Professor at the University of Chicago (with which she was associated since 1941), she made major discoveries regarding stereospecificity of enzymatic reactions. In the 60s, she started working on the photoinduced reactions in oxidative photosynthesis, i.e., in the Hill reaction (photoinduced O2 evolution in photosynthesis comes from water and not from CO2 and it evolves in two steps). She admired Otto Warburg (Nobel Prize in Physiology or Medicine in 1931), shared his methodologies and general ideas, and visited him several times in the Max Planck Institute for Cell Physiology in Berlin. Otto Warburg invited her to join the Institute as Director and she moved to Berlin in 1968.

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Birgit Vennesland

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After she scientifically criticized Warburg’s measurements of the quantum yield of photosynthesis, claiming that there had been a mistake in the procedure (perhaps too much nitrate in the medium), the situation was uncomfortable and the Max Planck Society created a special section called the Vennesland Section, physically separated from the Institute directed by Otto Warburg. She then studied, with her research group, the nitrate assimilation in photosynthetic organisms and made major contributions in this area [1]. She retired from the Max Planck Society in 1984 and went to live with her twin sister in Hawai where she continued teaching and working. She passed away in 2001. She advised several doctoral students and had many collaborators that continued the research in the areas she initiated. Birgit Vennesland received numerous awards and recognitions [2].

Eleanor Beatrice (Bea) Marcy Sweeney (Fig. 2). She was born in Boston (USA) in 1914. Since childhood, Beatrice was determined to be a scientist [3]. She showed enormous interest for nature around her. Sweeney was revolutionary for women in science. Her college botany professors, she explained, “…were fine women, but they were all spinsters without private lives.” She vowed to be different—to be a scientist and have a family. “After all, men in science did not have to give up family life.”

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Eleanor B. Marcy Sweeney

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After College, she raised 2 children while attending graduate school. In her autobiography [4], she recognized her graduate advisor who “…never in any way implied that I was inferior to his male graduate students. He quietly made it possible for me to continue my research during whatever time I could manage when I had my first child and then a second.”

She and her husband Paul Lee Sweeney had 4 children. In 1967, Sweeney joined the Biological Sciences Department at University of California Santa Barbara (UCSB), becoming a Professor in 1971 and an Emerita Professor in 1982. She immediately gravitated the College of Creative Studies (CCS) and began teaching in the new College, where she developed the “Walking Biology” course (a field research course), still being taught today. She served as CCS Associate Provost from 1978 to 1981. With the discovery of circadian rhythms in the luminescence of the dinoflagelate Lingulodinium polyedra (then called Gonyaulax polyedra), which she managed to culture in the laboratory, she opened a whole new area of research. Beatrice Sweeney is considered a pioneer researcher in circadian rhythms in plants and algae. The book she wrote on the subject, first published in 1969, was reedited in 1987 [5].

In the autobiography, written in 1987, she recommends: “I advise young women in science to keep their maiden names, dare to be assertive, and refuse to get discouraged. They must keep learning and reading, especially if they have to take leave from science for family reasons. It may not be easy to combine a scientific career with a family but it can be done and it’s worth all the effort. Research has sustained me through many otherwise unbearable times and given me much satisfaction, made my older years exciting” [4].

Bes Sweeney received various honors and awards and held several leading positions in learned societies. She mentored Carol Greyder (to be awarded the Nobel Prize in 2020) through her studying years at the CCS (UCSB). She died during a trip to a Gordon Conference in 1989 [6].

Germaine Stanier (Cohen-Bazire) (Fig. 3). Germaine Bazire was born in 1920 and educated in Toulouse, France. Germaine Bazire joined the laboratory of Jacques Monod (Nobel Prize in Physiology or Medicine in 1965) at the Pasteur Institute in 1950 and contributed actively to the heroic period of research on the mechanisms of enzyme induction. Her discovery, in 1953 with Jacques Monod, of the specific inhibition of the synthesis of tryptophan synthase by tryptophan, along with a similar and simultaneous observation on the specific inhibition of methionine synthase by methionine, led to the concept of end-product repression of biosynthetic pathways [7]. In 1953, Germaine discovered the world of the photosynthetic prokaryotes. In Berkeley, California, she then started research on the physiological, biochemical, and structural properties of the principal groups of photosynthetic prokaryotes: the purple and green anoxyphotobacteria and the cyanobacteria.

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Germaine Bazire

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Her research on cyanobacteria initiated in Berkeley and pursued after her return to Paris in 1971, and until her retirement, provided important results on the structure and function of phycobiliproteins in cyanobacteria [8]. Cohen-Bazire established herself as a world leader in the ultrastructure and physiology of cyanobacteria. She headed the Unité de Physiologie Microbienne, created by Stanier in 1971, from 1982 (after Stanier’s death) until her retirement in 1988, when it was named the Unité des Cyanobacteries [9]. Germaine Stanier (Cohen-Bazir) died in 2002.

Joan (Jan) M. Anderson (Fig. 4). She was born in 1932 in Queenstown, New Zealand. From early age, she was firmly determined to be a scientist and had to overcome serious difficulties, some related to her underprivileged social position [10]. She eventually obtained a BSc and a MSc from the University of New Zealand. She gained a fellowship to undertake postgraduate studies in the United States. Upon finding that the University of California, Berkeley, would not recognize her New Zealand degree, denying her access to the library, research facilities, and health insurance, she enrolled for a PhD at the UC Berkeley College of Chemistry (1956–59) under the supervision of Melvin Calvin (Nobel Prize in Chemistry in 1961) [11]. After successfully completing the degree and repaying the bond to New Zealand, she joined CSIRO (Commonwealth Scientific and Industrial Research Organisation) in Australia to work in the laboratory of Keith Boardman.

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Joan Anderson

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Anderson and Boardman managed to separate the two components of plant reaction centers (PSI and PSII) activated each by a different wavelength [12]. During a stay in Cambridge, she was forced by the circumstances (a coal’s miners strike that restricted electricity supply to the lab and a refrigerator explosion in the lab) to reflect on the research she had done up to that moment and wrote a review about the location of chlorophylls in the membrane, providing a model for the organization of photosynthetic membranes [13]. After the appearance of this review, many visitors started to pilgrim to CSIRO and collaborated with Joan Anderson. Fundamental discoveries on the structure of the photosynthetic membranes were done. A list of publications can be found at the repository [14]. Joan is widely recognized as one of the main contributors to the concepts we now have about the architecture of the photosynthetic membrane. She received many awards and was a mentor for many young female researchers. She held several positions in learned societies and served in the editorial boards of prestigious journals. Joan Anderson died in 2015.

Elizabeth Rohatsch Gantt (Fig. 5). She was born in 1934 in a small town near what is today the Serbian/Hungarian border, eventually moved with her family to the USA in 1949. She got her PhD in 1963 at Northwestern University [15]. After that she worked as a technician in Darmouth Medical School, but soon was engaged as a post-doctoral fellow with Samuel F. Conti and started working on the photosynthetic apparatus of red and green algae. They discovered in red algae the special complex between antennas and reaction centers and coined the name phycobilisomes [16]. She kept on working in this area at the Radiation Biology Laboratory of the Smithsonian Institution from 1967 until 1988. Then, she joined the Department of Botany at the University of Maryland, College Park (UMCP), where she became a professor and worked on several aspects of plant physiology, among others on the biosynthesis of carotenoids [17].

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Elizabeth Gantt

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In 1988, she was elected the first female president of the American Society of Plant Biologists. In 1996, she was elected member of the National Academy of Sciences. After the official retirement in 2007 as Distinguished Professor, she kept on participating in important projects [18]. Elizabeth Gantt was awarded the Darbaker Prize from the Botanical Society of America and the Gilbert Morgan Smith Medal of the National Academy of Sciences. The Dr. Elisabeth Rohatsch Gantt’58 Award was established by her Blackburn College class of 1958 classmates Carol (Nielson) Hughes and Clarence “Pete” Hughes as a way of recognizing and honoring Dr. Gantt’s great contributions to research in the field of plant science [19].

Helga Ninnemann (Fig. 6). She was born in 1938 in Königsberg (Ostpreussen in those days), emigrated at the end of World War II with her family to Brandenburg and later settled in Frankfurt/Main. During her thesis work in Frankfurt, she got fascinated with the light-regulated photomorphogenesis of liverworts [20]. She joined the lab of H. Stern and Warren Butler in La Jolla at the University of California, San Diego, as a postdoc (1966–1969), where she studied the effect of blue light on respiration in yeast [21] and started working on Neurospora. She continued working with Neurospora (among other subjects) until her last days. In 1979, she proposed that the flavin co-factor in nitrate reductase is responsible for the blue-light induction of conidiation and absorbance changes observed in Neurospora [22]. Much later, making a genetic analysis of the phenomenon, the photoreceptor for conidiation in Neurospora was identified as the FAD-(flavin adenine dinucleotide)-containing WC-1 [23].

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Helga Ninnemann

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During many years, she was interested in establishing pathogen-resistant potatoes by cell fusion techniques using resistant wild potato species. Together with her coworker Lieselotte Schilde, she succeeded in attracting several potato research grants. She investigated the involvement of phytoalexins in resistance and extended her research to alternative oxidase and mitochondrial function during pollen ripening in potato flowers. Several PhD students from South America joined her potato lab during this time [24].

Helga Ninnemann established a home in California and, after her return to Germany, she would frequently travel to California. Back in Tübingen, she joined the Institute of Chemical Plant Physiology, later renamed Institute of Plant Biochemistry. She was a great teacher and mentor, had many graduate students and went to the surrounding schools to propagate science at various levels. She encouraged many female students to join life sciences and to survive in the career path despite hard competition. She was an assiduous participant of the European Meeting on Photomorphogenesis, where I met her. Helga Ninnemann died in Tübingen in 2003 [24].

Daphne Vince-Prue (Fig. 7). Daphne Vince-Prue graduated from the University of London (Wye College) and did postgraduate work in plant physiology at UC Berkeley. She obtained her doctorates at the Universities of Reading and London, and subsequently taught plant physiology to students of horticulture and botany at Reading for most of her career. She has also been a Scientific Adviser and later Head of the Physiology and Chemistry Division at the Glasshouse Crops Research Institute for the Agricultural Research Council.

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Cover of the book “Science and the Garden: The Scientific Basis of Horticultural Practice”, by David S. Ingram (Editor), Daphne Vince-Prue (Editor), Peter J. Gregory (Editor)

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Since her retirement in 1986, she maintained contacts with research groups working in photoperiodism, and continued her interest in horticulture as a committee member of The Royal Horticultural Society. Her book “Photoperiodism in Plants” was published in 1975 and a 2nd edition was written together by Brian Thomas and published in 1996 by Academic Press. She was very active in the field photomorphogenesis [25].

The Book had its 3rd Edition in 2015 by Wiley-Blackwell (ISBN: 978-1-118-77843-2).

Helga Kasemir (Fig. 8). Helga was one of the several female biologists working in the research group of Hans Mohr at the Biology Institute of the Freiburg University in Germany. She worked on various aspects of phytochrome control of metabolic paths, such as chlorophyll biosynthesis [26, 27] as well as in other questions in plant physiology. She also collaborated with Hans Mohr after he took the Direction of the Akademie für Technikfolgenabschätzung (Academy for the Evaluation of the Consequences of Technology) in the German Province Baden-Württemberg, in 1995 [28].

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Helga Kasemir

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After her retirement in 2002, Helga changed gears, started painting and presented her paintings in several exhibitions.

Maria-Elizabeth Michel-Beyerle (Fig. 9). She was born in Germany in 1935, M-E. Michel-Beyerle studied Chemistry in Goettingen and Munich and got her PhD in the Technical University in Aachen. She got her Habilitation in Physical Chemistry in Munich while associated with H. Gerischer. She worked in several aspects of charge transfer in condensed phase. In 1976, she observed that upon quenching of singlet pyrene by N, N-diethylaniline in methanol at room temperature, the pyrene triplet and pyrene anion concentrations were modulated by a magnetic field [29]. This effect was interpreted as due to a magnetic field-dependent triplet production in the geminate radical ion pair, induced by hyperfine interaction of the unpaired electrons.

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M-E. Michel-Beyerle

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This was an unprecedented discovery and started the area of spin chemistry. The work on magnetic field-dependence of electron transfer processes was and is a very active area of research that led to important discoveries, for example analyzing primary processes in Photosynthesis. M-E. Michel-Beyerle started in 1981 a very successful Collaborative Research Centre (SFB) of the German Research Foundation (Deutsche Forschungsgemeinschaft) at the Technical University of Munich entitled “Elementary processes of photosynthesis”.

M-E. Michel-Beyerle was the speaker of this Center for many years and produced important contributions in the field of primary processes in photosynthesis, photovoltaic devices, and biosensors.

Since 2008, she has been a Visiting Professor at Nanyang Technological University in Singapore, and until 2021, she continued her research in the newly established “BioFemtoLabs” at that University. Together with R. A. Marcus (Nobel Prize in Chemistry in 1992), they produced important theoretical contributions on the mechanism of action of the UV-light plants photosensor UVR8 [30]. M-E. Michel-Beyerle is Professor Emeritus of Excellence at the Technical University of Munich and has received several awards [31].

Rosalie Crouch (Fig. 10). “A native of Norfolk, Virginia (USA), and a 1963 graduate of Randolph-Macon Woman's College, Rosalie K. Crouch, PhD, received her master’s degree in chemistry from Lehigh University in Pennsylvania in 1965 and her doctorate in organic chemistry from Albert Einstein University in New York in 1972” [32].

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Rosalie Crouch

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“Rosalie Crouch joined Medical University of South Carolina (MUSC) in 1975 as a faculty member in the College of Medicine’s departments of Ophthalmology and Biochemistry, and by 1982 had earned the title of professor. In 1989 Crouch was appointed acting dean of the College of Graduate Studies and became its dean in 1991. She held the title of dean concurrently with her 1995 appointment as the College of Medicine’s associate provost for research until 2000, at which time she became the Medical University’s first female provost and vice president for academic affairs.” Rosalie Crouch stepped down as provost and vice president for academic affairs in 2002 and returned to research.

Rosalie Crouch and her group have produced fundamental data regarding the visual cycle and about the molecular basis for certain ophthalmic disorders and illnesses [33, 34].

“In 2002 Rosalie was awarded the State of South Carolina Governor’s Award for Outstanding Service. She is a Distinguished University Professor of Ophthalmology and Biochemistry, and Provost Emerita. In 2003, Crouch was awarded the MUSC Distinguished Faculty Service Award, which recognized her for her 28 years of service, over 200 publications, 20-plus years of continuous NIH funding, and recognition by the National Eye Institute and Research to Prevent Blindness.”

In 2015, Rosalie was honored by the Medical University of South Carolina for her support to women faculty at MUSC, something Crouch had been doing for about 4 decades. During the award ceremony, she said: “[I am] trying to understand why there have not been more women at the top. Most seem to stop at the assistant or associate professor level” [32].

Silvia Elsa Braslavsky (the author) (Fig. 11). Silvia Elsa Braslavsky was born on April 5th, 1942, in Buenos Aires, Argentina, studied Chemistry in the school of Natural Sciences (University of Buenos Aires) and started her PhD studies in the same school in 1963 in the group led by Eduardo Lissi and Juan Grotewold. In October 1966, after the military intervention of the Argentinian Universities and the violent “Night of the Long Sticks” [35], she moved with the research group to Chile where she finished her doctoral work on gas phase kinetics. Her daughters Paula and Carolina were born in Chile in 1968 and in Buenos Aires, in 1969, respectively. After post-doctoral work at Penn State University (PSU), in 1972, she obtained an Associate Professor position at the newly created University of Rio Cuarto (Argentina), where she moved with her two daughters. At the end of 1974, the political situation strongly deteriorated, and she left, under extremely complex conditions, with her daughters again to PSU, and later to the University of Alberta in Edmonton, Canada. Already in Rio Cuarto, she had started thinking about working on phytochrome.

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Silvia E. Braslavsky

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This could be materialized in the Max Planck Institute for Radiation Chemistry in Mülheim an der Ruhr (Germany), where she went to work in October 1976, invited by the Institute Director, Kurt Schaffner. A productive work could be accomplished, first on open-chain tetrapyrroles as models for the phytochrome chromophore [36]. The spectroscopy of phytochrome extracted from etiolated oat seedlings was studied as well as of heterologously expressed plant phytochrome jointly with Wolfgang Gärtner. Studies by Alfred Holzwarth on ultrafast kinetics and by Peter Hildebrandt on resonance Raman spectroscopy, in the same Institute, led to a better knowledge of the details of the photocycle of phytochrome A (from etiolated plants) as well as of some other phytochromes [37, 38] Laser-induced optoacoustics, thermal lensing and photothermal beam deflection were applied for the understanding of the heat evolution after excitation of phytochrome, retinal proteins, and flavoproteins, considering that in these photosensors the quantum yield of photoconversion is either low (0.6 in retinal proteins) or very low in phytochrome (ca. 0.17), this meaning that a large amount of the absorbed energy is given back as heat (or as entropic changes as it turned out), of course subtracting fluorescence [39].

A ns-flash photolysis study of closed PSII reaction centers together with the recently available high resolution X-Ray structure of PSII led to the proposal of a photoprotection mechanism of the centers under high fluence conditions [40].

Since the recovery of Democracy in Argentina in 1983, Silvia interacted with research groups there and received several Argentinian post-doctoral collaborators in Germany. She also dedicated efforts to the work with the IUPAC Photochemistry Commission [41]. In 2007, she formally retired. Teaching a course on Biological Photoreceptors took her then to Ottawa, Barcelona, Havana, Santiago de Chile, Quito, Sao Paulo as well as to various Universities in Argentina.

The organization of the International Congress on Photobiology, which she chaired, held in 2014 in Córdoba, Argentina, was possible with the help of a newly created group of Argentine photobiologists (GRAFOB). Silvia is a Member of the Argentinian National Academy of Sciences (ANC), received the “Raíces” Prize from the Minister for Science and Technology (MINCyT, Argentina), the Finsen Medal of the International Union of Photobiology, the European Society of Photobiology medal, and is an International Ambassador of Photochemistry of the European Photochemistry Association (EPA).

Ana (Lorenzelli) Moore (Fig. 12). She was born in the Province of Buenos Aires (Argentina) on April 14th, 1942. Ana Lorenzelli studied Chemistry at the Universidad de La Plata (UNLP) and got a scholarship to go to Brazil for a master’s in chemistry, which she obtained in 1966 from the Universidade Federal do Rio de Janeiro. Later, she moved, together with her friends (also Chemistry students in the UNLP) Juana Chessa (Nita) Silber and Ernesto Silber, to Texas Tech University in Lubbock, where all three got their PhD degree. In Lubbock, Ana met Tom Moore in a Physical Chemistry course and they married. After a post-doctoral stay at the University of Washington, in 1976 Ana and Tom moved to Arizona State University (ASU) to the School of Molecular Sciences (formerly known as the Department of Chemistry and Biochemistry) where together with Devens Gust (who was at ASU since 1975) started a very successful team designing, synthesizing, and measuring the properties, of large and complex molecules that, upon irradiation, would mimic the photosynthesis process; thus, also called artificial photosynthetic constructs. The three of them were also founding members of one of the earliest centers at ASU, the Center for the Study of Early Events in Photosynthesis.

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Ana and Tom Moore in October 2018 in La Biela, a traditional café in Buenos Aires, together with figures of two legendary Argentinian writers: Jorge. L. Borges (left) and M. Bioy Casares (photo: SEB)

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The extraordinary knowledge and ability in organic chemistry Ana has, together with the physical chemistry abilities of Tom and Devens, constituted the basis for a very productive and innovative team that lasted as such until Devens Gust became Emeritus in 2014. The three co-authored  many of the papers coming out of the Centre and maintained collaborations with  a number of research groups worldwide. Their studies have advanced enormously the fundamental knowledge about light-induced electron transfer as well as, lately, on proton-coupled electron transfer (PCET) [42] and the molecular architectures and interactions that optimize these processes. The most cited paper with more than 2500 citations is Ref. [43].

Despite her enormous dedication, very important results, and excellence in teaching, it was not easy for Ana to get a position as professor. She was designated Associate Professor in 1989 (before that date she got a series of positions as Invited Professor, Research Assistant, Researcher), whereas Tom Moore was designated Assistant Professor already in 1976. Tom Moore became Professor in 1985 and Anna in 1996. “Having both members of a married couple in the same department was virtually unheard of”, said Gary Krahenbuhl, former Dean of the College of Liberal Arts and Sciences when celebrating 45 years of Ana Moore and Tom Moore at ASU, in 2022.

Ana Lorenzelli, Tom Moore, and Devens Gust became Regents Professors in 2011. Ana has kept collaborations with Argentinian researchers and has received (and still receives today) Argentinian students and post-doctoral fellows in the labs at ASU. Several of them are back in Argentina and have their own successful research groups. As a recognition, Ana (Lorenzelli) received in 2012 the “Raíces” Prize from the Argentinian Minister of Science and Technology (MINCyT) awarded to Argentinians working abroad and fruitfully collaborating with the Argentinian scientific community. Ana is still active, highly productive and offering beautiful lectures about their work.

Beverley Green (Fig. 13). Canadian Biochemist obtained her B.Sc. (Chemistry) from the University of British Columbia in 1960 and the PhD in Biochemistry from the University of Washington in 1965. Her major interest has been on chloroplasts. Her earlier research concerned the extended family of chlorophyll (Chl) a/b and Chl a/c light-harvesting proteins in plants and algae. Later, her group analyzed the structure of photosynthetic membrane proteins together with the evolution of the gene families that encode them, as well as the evolutionary origins of the chloroplasts themselves, investigating the minicircular genes found in dinoflagellate chloroplasts. These were the first chloroplast gene sequences ever reported for dinoflagellates, a large group of unicellular marine algae [44].

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Beverley Green

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Beverley Green is Professor Emeritus at the University of British Columbia, Canada.

Alison Telfer (Fig. 14). She was born in Great Britain in 1944, studied Botany at Oxford and moved to King’s College London for her PhD in 1966 working on the biochemistry of photosynthesis. Alison started working with Jim Barber in 1972 as a post-doctoral fellow [45]. She worked on photosynthesis, carotenes, chlorophylls, always associated with James (Jim) Barber (1940–2020) at Imperial College, financially supported by diverse research grants and could never obtain a permanent position, notwithstanding her many fundamental findings and creative activities. Alison Telfer remained with the group until Barber’s death. In 2004, Jim Barber wrote: …“I particularly wish to acknowledge my very long collaboration with Alison Telfer who joined my group as a post-doc in 1972 and who has survived 31 years of dealing with me” [46]. She was involved with the electrostatic stacking of thylakoid membranes and finally showed that when LHCII is phosphorylated under red light, this follows quenching of chlorophyll fluorescence as the phosphorylated-LHCII moves out of the stacked—Photosystem II (PSII) rich region of the chloroplast—to the unstacked areas where its energy goes to Photosystem I (PSI) and the two photosystems become more balanced [47].

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Alison Telfer

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Among several findings about the structure, behavior, and complexity of PSII, in 1994, Alison detected the formation of singlet molecular oxygen in isolated reaction centers of PSII and determined the formation quantum yield of this species [48]. Together with the group in the Technical University in Berlin, she characterized the hyperfine structure of 3P680 using pulse ENDOR spectroscopy [49]. Not finding the β-Car radical by ENDOR, Alison joined the lab of Bruno Robert at Saclay to look for Chl and Car radicals using Resonance Raman spectroscopy [50]. Alison Telfer was a member of a collective work on photosynthesis in a cyanobacterium grown in far-red light (750 nm) with the participation of chlorophylls absorbing in the red edge of the spectrum [51].

Alison Telfer is an Honorary Research Fellow at Imperial College, London (Great Britain).

Elaine M. Tobin (Fig. 15). Elaine M. Tobin was born in 1944 in Louisville, Kentucky (USA), she describes in an autobiography her early interest in nature, insects and biology in general and the difficulties she found to be able to follow science courses as a girl and to be taken seriously in those courses [52]. She also describes the support she got from some of her scientific guides, especially from Winslow Briggs, her PhD thesis supervisor [53]. In fact, before joining Briggs’ lab, she intended to apply for a PhD position at the Biochemistry Department in Stanford University, but the Department did not accept women in their PhD courses, “because women do not get Nobel Prizes” [54]. After attending Winslow Briggs’ Plant Physiology course, Winslow accepted her in his lab and eventually Elaine moved as a graduate student to Harvard with Winslow Briggs, when he accepted a professor position there.

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Elaine Tobin

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After a year at the Weizmann Institute in Israel, and a post-doctoral stay at Brandeis, in 1975, she established herself (not without difficulties as a woman) in the University of California (Los Angeles, USA) where she and her collaborators have produced important pieces of work such as the first proof that phytochrome action could affect transcription of specific genes [55].

Tobin’s experiments provided proofs, at the same time as the lab from G. Coupland, that some genes (in her case the circadian clock associated gene, CCA-1, and LHY in Coupland’s lab) are components of the central oscillator in plants [56, 57].

Elaine was the organizer and chair of the 1985 Gordon Conference on Plant Physiology. She was a member of the Editorial Board of several Journals and member of important research granting panels. Elaine Tobin retired in 2014. In the latest publications from her lab, the focus remained on the regulation of the circadian clock in plants.

Nicole Tandeiau de Marsac. A French biologist born in 1944, made major contributions in the areas of cyanobacteria, phycobilisomes and phycobiliproteins. She did fundamental work on evolution of cyanobacteria, including interesting contributions to the knowledge on phytochromes [58]. She studied the phenomenon of chromatic adaptation in cyanobacteria. Nicole co-authored several papers with Germaine Stanier (Cohen-Bazir) (see above). One of Nicole’s latest papers was on the revision of the genus Planktothrix [59]. Another aspect of Nicole’s interest was the cyanobacteria of sweet waters and their toxicity. Her group studied the genome of the highly toxic Microcystis aeruginosa and identified the regions coding for the toxic metabolites [60]. Other genomes of cyanobacteria were studied in her laboratory.

Nicole was the Head of the Cyanobacteria collection in the Unit of Microbial Physiology at the Institut Pasteur (Paris) until her retirement in 2009. Nicole passed away in the Provence on July 30, 2020.

Joan Roberts (Fig. 16). Joan Roberts was born on October 23rd, 1944. Joan is Professor of Chemistry at Fordham University. Although trained as a classical Organic Chemist (synthesis and mechanisms), her main area of research has been to study the photophysics, photochemistry and photobiology of natural and synthetic (drugs, nanoparticles) chromophores in the human eye [61, 62]. Her initial ocular studies were defining the mechanism(s) of UV radiation interactions with Xanthurenic Acid (natural), and photodynamic dyes and drugs (synthetic) that might induce cataracts. These studies were extended to examine how short-wavelength blue light might interact with retinal chromophores that would lead to macular degeneration.

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Joan Roberts

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Joan worked on improvement of photodynamic therapy (PDT) against cancer (including ocular melanoma), using selective protection of normal tissues from PDT side effects [63, 64]. “We have contributed to the understanding of how visible light directed through the eye can disrupt both sleep and the immune response (photoneuroimmunology) by interfering with the synthesis and chemistry of melatonin, which is not only synthesized in the pineal gland but is also synthesized by retinal photoreceptors” [65,66,67]. “None of this interdisciplinary research could have been accomplished without the collaboration with fellow scientists, especially Dr. Dan-Ning Hu, those from ESP and ASP and former post-doctoral fellows Zhao Baozhong and Albert Wielgus.” Joan Roberts received in 2017 the medal of the European Society of Photobiology for “Excellence in Photobiological Research”.

Tiina Karu (Fig. 17). Tiina Karu was born on April 19th, 1947, in Tartu (Estonia). She received her master’s degree in physics also in Tartu in 1970, and obtained her PhD in Moscow in 1974 and the title of Doctor of Science in Physics in St. Petersburg in 1990. Since 1980, she is head of the laboratory in the Laser Technology Research Center of the Russian Academy of Sciences in Troitsk, Russia. She has focused her research on the effects of low power, low energy irradiation on cells and living organisms. She proposed, after experimenting with cells, that cytochrome in the mitochondria is the absorbing compound responsible for the effects of low intensity red light in some treatments [68].

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Tiina Karu with Kendrick Smith (Stanford University) in a Laser Conference in Tokyo in 2009; photo: Kendrick Smith Webpage

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Already in 1989, Tiina Karu published the book “Photobiology of low-power laser therapy” (Routledge) and she said: “This book provides a concise review of the efforts undertaken to demonstrate or disprove the existence of photostimulation by low-power lasers. Collating the factual evidence gathered so far, this volume contains extensive discussions of the action (with possible mechanisms) of monochromatic visible light on cells and organisms and of primary photoreceptors. Other topics such as short-term responses to irradiation are also included.”

In 1998, CRC Press published another book authored by Tiina Karu: “Science of Low Power Laser Therapy” (ISBN-13: 978-90-5699-108-1, ISBN: 90-5699-108-6). Her unbiased experiments have given scientific basis to the low-power red-light therapy, in particular the method called photobiomodulation [69].

Since 1990, Tiina Karu is Professor at the Russian Academy of Laser Sciences and in 1998 was made an Academician in the Russian Academy of Laser Sciences.

Eva-Mari Aro (Fig. 18). She was born in Finland in 1950, received her master’s in 1975 and her PhD degree in 1982 from the University of Turku in Finland. She was appointed as Associate Professor at the same University and became full Professor of Plant Molecular Biology in the Department of Biochemistry in 1992. E-M. Aro has made major contributions to the knowledge on photosystem II (PSII), its damage and recovery as well as the acclimation to changing environmental conditions. One of her contributions with ca. 2000 citations is a paper on the stability and damage recovery of PSII [70]. Her research group has produced important contributions to the biochemical changes in the chloroplasts induced by different light fluences [71].

Fig. 18
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Eva-Maria Aro

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Aro’s more recent research interests are applied bioenergy research and synthetic biology approaches, i.e., applying photosynthetic principles towards production of sustainable fuels and chemicals in cyanobacteria cell factories.

Eva-Mari Aro received the honorary title of Academician of Science in 2017 and was awarded a Commander First Class of the Order of the Lion of Finland in 2019. In 2018, Aro was elected as foreign fellow to the US National Academy of Sciences. She is a member of the Finnish Technical Sciences Academy (2012) and the Finnish Academy of Science and Letters (2002), and an honorary doctor of University of Helsinki (2014) and Umeå University (2012) as well as an honorary professor of the Chinese Academy of Science (2006). Aro has chaired two Academy of Finland Centres of Excellence and the NordAqua project with nine other partners from Nordic counties. She also served the Finnish Academy of Science and Letters 2010–2015, first as a board member, then as a vice president and president, and acted during 2004–2007 as president of the International Society of Photosynthesis Research. She was a founding partner of the EU funded CSA initiative “SUNRISE: Solar Energy for a Circular Economy” (https://sunriseaction.com/, EU R&T Roadmap 2019) and currently serves as a Finnish contact for SunERGY EU CSA (https://www.sunergy-initiative.eu), promoting emerging biotechnologies for industrial applications.

Marta Cecilia del Carmen Bunster Balocchi (Marta Bunster) (Fig. 19). Born in Concepción (Chile), on May 27th, 1951, she studied Biochemistry at the University of Concepción, where she graduated with a work on X-rays diffraction of synthetic polypeptides [72]. After a short stay in Santiago (Chile), she returned to Concepción where she obtained a PhD degree in 1981 working partially at the University of Florida (USA). Back in Concepción, in 1981, she met Hilda Cid, a Chilean crystallographer who had left the Country together with her family in 1973, due to political persecution after the military putsch and was back from Sweden with equipment for X-rays crystallography provided by the University of Uppsala. Marta Bunster and Hilda Cid established the Molecular Biophysics Laboratory at the Faculty of Biological Sciences and Natural Resources, now the Faculty of Biological Sciences, in the University of Concepción. Marta Bunster investigated phycobilisomes [73, 74], and several other proteins such as proteases from Antarctic bacteria (technologically useful cold-adapted enzymes) [75], with the aim of understanding the energy transfer mechanisms operating between the components of the phycobilisomes vectorially into the reaction center.

Fig. 19
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Marta Bunster

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In the early 2000s, Marta Bunster dedicated major efforts to consolidate international cooperation in bioinformatics; in 2002, she was a founding member of the Iberoamerican Network for Bioinformatics, later renamed as Iberoamerican Society for Bioinformatics (SoIBio). Marta Bunster was the Secretary in the first executive board of SoIBio and remains active in the board in 2023. In December 2022, Marta Bunster received the Emeritus Award from the Chilean Society of Biochemistry and Molecular Biology for her trajectory of research and teaching [76].

Marie-Michèle Cordonniere-Pratt (Fig. 20). She was born on September 28th, 1951, in Haute-Savoie (France) and got her PhD in cellular biology in 1977 from the University of Rouen, France. She continued in 1987 with a French “Doctorat d’Etat” from the University of Paris 6. After spending a short postdoc in the lab of Lee Pratt at the University of Georgia in the late 70s, she went to work for a full year at the US Department of Agriculture in Beltsville, establishing there a new laboratory equipped for research in phytochrome (bilin-binding photoreceptor found in higher and lower plants, fungi, and photosynthetic and non-photosynthetic bacteria, as we know today). In 1980, she received a junior faculty position at the University of Geneva and expanded her immunochemical work there with independent support from the Swiss National Funds. In 1985, she was recruited by Ciba Geigy (now part of Novartis) to create a new immunochemically oriented laboratory within an equally new plant-oriented research facility in Research Triangle Park, North Carolina.

Fig. 20
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Marie-Michèle, photo taken by Lee Pratt, 2023. Mont Blanc in the back

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Marie-Michèle was possibly the first to create monoclonal antibodies for plant research (having spent time in Basel and Zurich to learn the technology in the late 70s) and her achievement led to numerous expansions of the use of monoclonal antibodies (immunodetection, immunocytochemistry, immunopurification, etc.) in a non-medical research laboratory. She was certainly a pioneer in the production of antibodies for phytochrome [77], also to differentiate the red absorbing form of plant phytochrome (Pr) from the far-red absorbing form (Pfr) [78]. Marie-Michèle moved to the University of Georgia (US), together with Lee Pratt (they got married in 1990), where she continued a very productive work on the antibodies for phytochrome (s), especially for the identification of the various types of the plant photosensor [79]. Together they started a lab on Genomics and Bioinformatics at the University of Georgia (UGA) developing informatic tools for the analysis of evolutionary paths, also in the phytochrome family [80], and extended it to other organisms. For almost a decade in the 1990s both Marie-Michèle and Lee Pratt served as foreign research advisors to a laboratory at the Riken Institute of Japan in Wako, Japan.

Marie-Michèle and Lee became Emeriti at UGA in 2007 and turned towards other activities, going on expeditions to remote places, discovering and enjoying native populations and learning more about primitive/wild environments.

María Vernet (Fig. 21). María Vernet was born in the austral autumn (March 28th) of 1952 in Buenos Aires, Argentina, as the second in a family of five siblings. Her first hiking trip to Cerro Lopez in the Andes, near San Carlos de Bariloche, when she was 6 years old, helped her discover a love of the outdoors that translated later into a career in Earth and Biological Sciences. After attending a high school concentrating in the humanities, Maria trained as an ecologist at the University of Buenos Aires, where she received her Licenciatura in Biology (between Bachelor and Master in Science). During that time, a fellowship in marine phytoplankton, studying diatoms, connected her with the mystery and beauty of marine life. Her first job in the Patagonia coast helped her discover that scallops' food consisted of diatoms living in marine sediments, small micro-algae that become suspended in ocean water by tidal currents. The need to specialize in oceanography took her to Seattle, USA, where she obtained a master in science and a PhD in Oceanography at the University of Washington. At the time of the discovery of the ozone hole over Antarctica, her first project as a post-doctoral researcher investigated the effect of increased ultraviolet light reaching the ocean. A life-long career in the photobiology of polar oceans was thus born.

Fig. 21
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Maria Vernet

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Sunlight reaching the ocean surface is the driver of marine photosynthesis, responsible for half the annual oxygen input to the atmosphere. Both incident light and its transmission into ocean waters define the underwater light field, where longer wavelengths in the red are rapidly absorbed (first meter) and the blue wavelengths are transmitted the deepest. Thus, defining not only irradiance but light quality are essential measurements in marine photobiology. María’s PhD thesis concentrated in identifying and quantifying photosynthetic pigments in phytoplankton and their role in light absorption.

Photosynthetic pigments have evolved with algal diversification such as they can be used as taxonomic indicators of difficult to identify micro-algae. María’s PhD thesis in 1983 was the first to quantify an unknown pigment in tropical oceans later identified as chlorophyll a2, belonging to a new group of marine cyanobacteria, Prochlorococcus [81]. Her publications in the late 1980s and early 1990s were among the first to show photoinhibition of photosynthesis by underwater ultraviolet radiation, mainly UV-B [82]. A continent-wide project from Canada to the Antarctic Peninsula, with latitudinal changes in ultraviolet irradiance and quality, with international collaborators from 5 countries, and associates from several Latin American countries, showed for the first time the complexity of the effect of ultraviolet radiation on aquatic ecosystems and the variety of strategies in plant and animal life to avoid, adapt or mitigate harmful radiation [83]. During the 1990s and 2000s, as an assistant, associate and then full researcher at Scripps Institution of Oceanography in California, USA, her studies in photobiology concentrated on spatial and temporal variability of marine photosynthesis in Antarctic coastal waters due to the role of the environment in shaping light irradiance, such as light transmission through sea ice and the annual demise and initiation of photosynthesis at the beginning and end of the polar night. Those studies discovered that the amount and distribution of winter sea ice affected the irradiance necessary to kick-off phytoplankton growth during the austral spring and how it shaped the timing of iron limitation later in the season (note the nowadays dramatic shrinking of arctic and Antarctic sea ice, and the earlier start of phytoplankton and seawater warming). In the following decade, environmental photobiology became part of understanding global climate change, where sunlight is part of a complex interplay of multi-stressors that add up, subtract or multiply their effects, both positive or negative, on polar marine photosynthesis.

In the last few years, as Emerita researcher, María has developed a citizen science project in the Antarctic Peninsula, working with IAATO, the international association of tourist ships, to disseminate her love of polar photobiology to non-scientists and improve sampling opportunities to graduate students and young scientists [84].

Diana Kirilovsky (Fig. 22). Born as Diana Tiffenberg in Buenos Aires on October 21st, 1954, she left the country in 1976 at a very difficult and dangerous time in Argentina, after the military coup of Rafael Videla and his accomplices. Diana Tiffenberg-Kirilovsky went to Israel together with her husband. Without family and friends, in exile, they first had to learn Hebrew, a language totally unknown for them. In 1981, when she was starting her work for the PhD thesis with I. Ohad, their son was born and in 1984, their daughter followed. She obtained her Ph.D. in Biochemistry at the Hebrew University of Jerusalem in 1986, working on phycobilisomes.

Fig. 22
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Diana Kirilovsky

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The family moved to France to a new Country with a new language for a post-doctoral position at CNRS and CEA laboratories. Life was not easy and recognition came later than for her male colleagues. In 1991, she obtained a Senior Research Scientist position at the CNRS, and she worked in several laboratories in Gif sur Yvette, L’ENS and CEA Saclay. Diana Kirilovsky has studied the molecular biology and physiology of cyanobacteria for over 35 years focusing on the role of light as a source of stress and as a regulator.

Diana studied the mechanisms of photoinhibition, photoprotection and light acclimation in cyanobacteria. A major contribution of her group was the discovery and characterization of a new mechanism of photoprotection in cyanobacteria involving a unique type of photoactive carotenoid protein, the Orange Carotenoid Protein (OCP). OCP is a blue-light photoreceptor sensing light intensity which has a completely different photocycle than other photoreceptors [85]. So far, OCP is the first photoactive protein containing a carotenoid as active chromophore. OCP is also an excellent model protein to study the protein influence on the electronic properties of carotenoids, considering that the carotenoid functions in photoprotection, in vision and as antioxidant, are tightly bound to their electronic properties [86].

Diana Kirilovsky was a member of the committees of the French and International Photosynthesis Society during several years and of the French Photobiology Society. She retired as Research Director “Classe Exceptionnel” (DRCE1, CNRS) and as Group Leader in the Institute of Integral Biology of the Cell (I2BC, CNRS). Diana Kirilovsky has been an active participant of the Group of Argentinian Molecular Photobiologists (GRAFOB for its name in Spanish) and has taken part in many of the meetings of GRAFOB since the group was created in 2011. Diana Kirilovsky is Emeritus Research Director in the Institute of Integral Biology of the Cell (CNRS).

Joanne Chory (Fig. 23) [87]. Joanne Chory was born in Boston, Massachusetts (USA) in 1955. She graduated at Oberlin College in Ohio with a degree in biology with honors. She received her PhD degree from the University of Illinois Urbana-Champaign. Considered the most influential plant biologist of the modern era and one of the greatest scientific innovators of our time, Chory’s 30 years of work has pioneered the use of molecular genetics to study how plants change their shape and size to optimize photosynthesis and growth for different environments. Joanne Chory is a Professor and Director of the Plant Molecular and Cellular Biology Laboratory, at the Salk Institute for Biological Studies and an investigator of the Howard Hughes Medical Institute. With her group she has made important contributions to the understanding of light effects in plants. In 1994, they proposed that the DET1 gene product is the protein that integrates the light signals in plants [88]. In her lab, the role of brassinosteroids as plant hormones was discovered [89]. The latter publication received almost 1200 citations.

Fig. 23
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Joanne Chory

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In a multiple lab effort under Joanne Chory’s leadership, the mechanism of biosynthesis of auxin, that triggers the effects known as shade avoidance in plants, was disentangled [90].

The latest big project in Joanne Chory lab is the Harnessing Plants Initiative, a project made up of five teams of plant biologists at Salk that are attempting to fight climate change by engineering plants to take in more carbon dioxide through their roots [91]. For this project she has obtained large financial support.

On a personal side, Joan and her husband have adopted four children.

Joanne Chory has received numerous awards. Since 1999 she is a member of the U. S. National Academy of Sciences, in 2006 became Associate Member of EMBO, in 2008 was elected member of the German Academy of Sciences Leopoldina, in 2009 was elected Foreign Associate of the Académie des Sciences, France, in 2011 was elected Foreign Member of the Royal Society, London, and in 2019 received the Princess of Asturias Award for Technical and Scientific Research.

1 Final words

The short biographical sketches of the women above show us very diverse examples of hard work, creative approaches, and strength to overcome difficulties, often due to their being a woman. Looking at the general situation in the academic world, there is no doubt that changes have occurred, even a couple of women have being awarded the Nobel Prize after Marie Curie in 1903 and 1911, and Irene Joliot-Curie in 1935; in Physics: Maria Goeppert Maier in 1963, Donna Strickland in 2018, Andrea Ghez in 2020 and Anne L'Huillier in 2023; in Chemistry: Dorothy Crowfoot Hodgkin in 1964, Ada Yonath in 2009, Frances Arnold in 2018, Jennifer Doudna and Emmanuelle Charpentier in 2020, and Caroline Bertozzi in 2022. 14 women got the Nobel Prize in Physiology or Medicine (https://www.nobelprize.org/prizes/lists/nobel-prize-awarded-women/).

Many women born after 1955 are presently doing very interesting and diverse research on photoreceptors in many parts of the world. Here is a list of those I have identified, in alphabetical order of their surname: Margaret Ahmad (France), Virginia Albarracín (Argentina), Cornelia Böhm (Austria, Finnland), Claudia Büchel (Germany), Paula Casati (Argentina), Roberta Croce (The Netherlands), Cristina Dorador (Chile), María Eugenia Farias (Argentina), Katrina Forest (USA), Petra Fromme (USA), Christiane Gatz (Germany), Leticia González (Austria), Elena Govorunova (USA), Muriel Gugger (France), Karen Halliday (UK), Joanna Kargul (Poland), Cyril Kerfeld (USA), M. Gabriela Lagorio (Argentina), Aba Losi (Italy), Sarah Mathews (Australia), Benedetta Mennucci (Italy), Sabeea Merchant (USA), Maria Mittag (Germany), María Andrea Mroginski (Germany), María A. Mussi (Argentina), Elena Orellano (Argentina), Laura Pedraza González (Italy), Jimena Rinaldi (Argentina), Ruth Rosenstein (Argentina), Ilme Schlichting (Germany), Emina Stojkovic (USA), Annegret Wilde (Germany), Eva Wolf (Germany), and Atina Zouni (Germany). The latter is a biased, partial collection of names of female scientists who are presently shaping the knowledge, together with their male colleagues, in the wide area of biological photoreceptors.

Presently, the number of women in leading administrative positions in the academic world in North America and in the European countries is increasing. In Argentina, there have been many women participating in the academic world already since several decades. Yet, very few women in Argentina have acted as School Deans or University Rectors (no woman has ever been elected as a Rector at the leading University of Buenos Aires) or have reached the highest level in the scale of the National Research Council (CONICET for its name in Spanish). The number of women in the editorial boards of the journals as well as in the executive board of the learned societies and in the Academies has increased in the last years. The current President of the Argentinian Research Council is Ana Franchi, a female biological chemist.

Notwithstanding the improvements, a general look at the numbers in the world tells us that there is still a lot to improve. The future is bright, but it is necessary to strive for it.