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Ruth Lyttle Satter (March 8, 1923 – August 3, 1989) was an American botanist best known for her work on
circadian A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., endogenous) and responds to ...
leaf movement.


Biography

Ruth Lyttle Satter was born March 8, 1923, in
New York City New York, often called New York City or NYC, is the most populous city in the United States. With a 2020 population of 8,804,190 distributed over , New York City is also the most densely populated major city in the U ...
as Ruth Lyttle.Arthur W. Galston: "RUTH LYTTLE SATTER March 8, 1923–August 3, 1989", J Biol Rhythms March 1991 6: 2–3,
bibliographic detailspreview
/ref> Satter received a B.A. in mathematics and physics from
Barnard College Barnard College of Columbia University is a private women's liberal arts college in the borough of Manhattan in New York City. It was founded in 1889 by a group of women led by young student activist Annie Nathan Meyer, who petitioned Col ...
in 1944. After graduating, she worked at
Bell Laboratories Nokia Bell Labs, originally named Bell Telephone Laboratories (1925–1984), then AT&T Bell Laboratories (1984–1996) and Bell Labs Innovations (1996–2007), is an American industrial research and scientific development company owned by mul ...
and Maxson Company. In 1946 she married Robert Satter and in 1947 she became a homemaker, devoting herself to raising her and Robert's four children, Mimi, Shoshana, Jane and Dick. While raising her children, her love of plants led her to complete the
New York Botanical Garden The New York Botanical Garden (NYBG) is a botanical garden at Bronx Park in the Bronx, New York City. Established in 1891, it is located on a site that contains a landscape with over one million living plants; the Enid A. Haupt Conservatory, a ...
’s horticulturist training in 1951 and to serve as a horticulture instructor for the YMCA Hobby School from 1953 to 1963. In 1964, she began her graduate studies in
plant physiology Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants. Closely related fields include plant morphology (structure of plants), plant ecology (interactions with the environment), phytochemistry (bi ...
at the
University of Connecticut The University of Connecticut (UConn) is a public land-grant research university in Storrs, Connecticut, a village in the town of Mansfield. The primary 4,400-acre (17.8 km2) campus is in Storrs, approximately a half hour's drive from H ...
, where she earned her PhD in botany in 1968. During her doctoral work, Satter began unraveling the molecular underpinnings of the plant circadian clock. She determined the impact of red/far red light and the associated photopigment, phytochrome, on plant
morphogenesis Morphogenesis (from the Greek ''morphê'' shape and ''genesis'' creation, literally "the generation of form") is the biological process that causes a cell, tissue or organism to develop its shape. It is one of three fundamental aspects of deve ...
. Her work on
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., endogenous) and responds to ...
s would define her career and markedly impact the field of
chronobiology Chronobiology is a field of biology that examines Time, timing processes, including periodic (cyclic) phenomena in living organisms, such as their adaptation to sun, solar- and moon, lunar-related rhythms. These cycles are known as biological rh ...
. In 1968, after completion of her PhD, Satter joined the lab of Arthur W. Galston at
Yale University Yale University is a Private university, private research university in New Haven, Connecticut. Established in 1701 as the Collegiate School, it is the List of Colonial Colleges, third-oldest institution of higher education in the United Sta ...
to work first as a staff biologist and then as a research associate. At Yale, Satter continued her research on plant chronobiology by studying control of leaf movements. Her work demonstrated that ion flux in leaf motor cells drives this movement, and that the same mechanisms control the movement if the plant is in an environment with a light-dark cycle or an environment with constant light or constant darkness. Additionally, in 1980, she co-authored the third edition of ''The Life of the Green Plant'', a textbook on plant physiology and botany, with Galston and another colleague, Peter J. Davies.Galston, A. W., Davies, P. J., & Satter, R. L. (1980). The life of the green plant. Prentice-Hall ; hbk ; pbk In 1980, Satter also became a professor-in-residence at the
University of Connecticut The University of Connecticut (UConn) is a public land-grant research university in Storrs, Connecticut, a village in the town of Mansfield. The primary 4,400-acre (17.8 km2) campus is in Storrs, approximately a half hour's drive from H ...
, where she discovered that the
phosphatidylinositol Phosphatidylinositol (or Inositol Phospholipid) consists of a family of lipids as illustrated on the right, where red is x, blue is y, and black is z, in the context of independent variation, a class of the phosphatidylglycerides. In such molecul ...
cycle is the basic light transduction mechanism in the leaf motor cells. The same year Ruth was diagnosed with
chronic lymphocytic leukemia Chronic lymphocytic leukemia (CLL) is a type of cancer in which the bone marrow makes too many lymphocytes (a type of white blood cell). Early on, there are typically no symptoms. Later, non-painful lymph node swelling, feeling tired, fever, n ...
but this diagnosis only heightened her scientific focus. During this time she published multiple papers, worked on a book, and gained international acclaim for her work on chronobiology. She also stayed active with her husband as they traveled the world to bike, swim, and experience different cultures.


Death and legacy

While Satter had remained active throughout most of her illness, her health began to deteriorate in the late 1980s. As her quality of life plummeted, she chose to end treatment and take control of her remaining time. Her peace with this decision and her final days were immortalized by her husband in a
New York Times ''The New York Times'' (''the Times'', ''NYT'', or the Gray Lady) is a daily newspaper based in New York City with a worldwide readership reported in 2020 to comprise a declining 840,000 paid print subscribers, and a growing 6 million paid ...
article. Satter finally lost her protracted battle with leukemia at the age of 66 on August 3, 1989. Satter's will established the Ruth Satter Memorial Award which gives $1000 to women who interrupted their education for at least three years to raise a family.Austin, R. (1997). The Grants Register 1997. New York: Macmillan In 1990, the
Ruth Lyttle Satter Prize in Mathematics The Ruth Lyttle Satter Prize in Mathematics, also called the Satter Prize, is one of twenty-one prizes given out by the American Mathematical Society (AMS). It is presented biennially in recognition of an outstanding contribution to mathematics ...
was also established in her memory, with funds donated by her sister, Joan S. Birman.AMS::Ruth Lyttle Satter Prize in Mathematics. (n.d.). Retrieved April 23, 2019, from http://www.ams.org/prizes-awards/paview.cgi?parent_id=35 The prize is given every two years and honors Satter's commitment to research and encourages women in science by awarding $5000 to a woman who has made a significant contribution to mathematics research.


Contributions to chronobiology


Mechanisms of circadian leaf movement

Circadian leaf movement was first observed by the French monk
Jean-Jacques d'Ortous de Mairan Jean-Jacques d'Ortous de Mairan (26 November 1678 – 20 February 1771) was a French geophysicist, astronomer and most notably, chronobiologist, was born in the town of Béziers on 26 November 1678. De Mairan lost his father, François d'Ortous ...
in the 18th century. It was such a critical method for observing circadian rhythms in plants that the phenomena was dubbed the "hands of the circadian clock". However, it was not until Satter's groundbreaking work that the molecular mechanisms of this phenomenon were understood. Satter worked to decipher the structure and function of
pulvini A pulvinus (pl. ''pulvini'') is a joint-like thickening at the base of a plant leaf or leaflet that facilitates growth-independent movement. Pulvini are common, for example, in members of the bean family Fabaceae (Leguminosae) and the prayer plan ...
, specialized plant motor organs found at the base of leaves and leaflets which allow leaves to extend and fold down. She identified that changes in
potassium Potassium is the chemical element with the symbol K (from Neo-Latin '' kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosp ...
and
chloride The chloride ion is the anion (negatively charged ion) Cl−. It is formed when the element chlorine (a halogen) gains an electron or when a compound such as hydrogen chloride is dissolved in water or other polar solvents. Chloride s ...
ion concentrations in pulvini drive
osmotic Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region of ...
water flux, or changes in the amount of water in the cell. The pulvini contain two cell types, flexor and extensor cells. Satter's work demonstrated that as the flexor gains solutes and thus water to increase their rigidity, the extensor loses ions and water to decrease their rigidity. The coordinated inverse rigidity changes in extensor and flexor induce extension or collapse of the pulvinus to lift and lower leaflets. Satter also collaborated with Richard Racusen of the
University of Vermont The University of Vermont (UVM), officially the University of Vermont and State Agricultural College, is a public land-grant research university in Burlington, Vermont. It was founded in 1791 and is among the oldest universities in the United ...
to study circadian changes in pulvini
membrane potential Membrane potential (also transmembrane potential or membrane voltage) is the difference in electric potential between the interior and the exterior of a biological cell. That is, there is a difference in the energy required for electric charge ...
. Satter determined that the changes in membrane potential were too quick to be explained by the passive movement of potassium ions reported in her earlier papers. She discovered an energy-consuming
proton pump A proton pump is an integral membrane protein pump that builds up a proton gradient across a biological membrane. Proton pumps catalyze the following reaction: :n one side of a biological membrane/sub> + energy n the other side of the membra ...
that removes protons (H+) from the cell, facilitating the observed rapid electrical changes as well as potassium ion flux. Thus, Satter and Racusen determined that changes in membrane potential drives the ion flux necessary for circadian leaf movement. Additional experiments observing the pH in both flexor and extensor pulvini cells in different experimental conditions (darkness, white light, red light, and far red light) demonstrated that pH levels in flexor and extensor cells were inversely correlated. From these studies, Satter demonstrated that proton pumps were active in flexor cells in darkness and in extensor cells in light.


Entrainment of motor cells

Satter additionally unveiled the mechanisms that allow circadian leaf movements to synchronize with (entrain to) light-dark cycles. She showed that
phytochrome Phytochromes are a class of photoreceptor in plants, bacteria and fungi used to detect light. They are sensitive to light in the red and far-red region of the visible spectrum and can be classed as either Type I, which are activated by far-re ...
s, a type of plant
photopigment Photopigments are unstable pigments that undergo a chemical change when they absorb light. The term is generally applied to the non-protein chromophore moiety of photosensitive chromoproteins, such as the pigments involved in photosynthesis and pho ...
, mediate changes in membrane potential of pulvini in response to red and far-red light.Sage, L. C. (1992). Pigment of the Imagination: A History of Phytochrome Research. Burlington: Elsevier Science. Phytochromes have two interconvertible
isomeric In chemistry, isomers are molecules or polyatomic ions with identical molecular formulae – that is, same number of atoms of each element (chemistry), element – but distinct arrangements of atoms in space. Isomerism is existence or possibil ...
forms, Pfr and Pr. Red light converts the pulvinus phytochrome to the Pfr form which promotes leaf closing, while exposure to far-red light converts the photopigment to the Pr form and promotes leaf opening. Pfr phytochrome depolarizes the cell membrane by opening potassium
ion channel Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by gating the flow of ...
s and facilitating potassium ion flow while Pr hyperpolarizes the cell membrane to close the potassium ion channel and prevent ion flow. Over the course of 24 hours, the amount of red and far-red light in the environment changes, so the form of phytochrome in the pulvinus also changes over time, causing the plant leaves to open or close as a result of the environmental light cycle. Satter also studied other methods of leaf movement entrainment including the effects of blue light. Satter and her colleagues showed that blue light leads to
phase shifts In physics and mathematics, the phase of a periodic function F of some real variable t (such as time) is an angle-like quantity representing the fraction of the cycle covered up to t. It is denoted \phi(t) and expressed in such a scale that it v ...
in leaf movement in both ''
Samanea saman ''Samanea saman'', also sometimes known as the rain tree, is a species of flowering tree in the pea family, Fabaceae, now in the Mimosoid clade and is native to Central and South America. Its range extends from Mexico south to Peru and Braz ...
'' and ''
Albizia ''Albizia'' is a genus of more than 160 species of mostly fast-growing subtropical and tropical trees and shrubs in the subfamily Mimosoideae of the family Fabaceae. The genus is pantropical, occurring in Asia, Africa, Madagascar, America an ...
''. Satter found that blue light exposure can affect when plant leaves extend and that the timing of blue light exposure determined whether leaves extended earlier or later than expected. While these studies revealed that blue light could advance or delay circadian leaf movement rhythms, the photoreceptor that mediates this response would not be discovered until later.


References


External links


1988 interview with Satter
{{DEFAULTSORT:Satter, Ruth Lyttle Chronobiologists Plant physiologists 1923 births 1989 deaths American women biologists American women botanists American women mathematicians Barnard College alumni University of Connecticut alumni University of Connecticut faculty Deaths from leukemia 20th-century American botanists 20th-century American mathematicians 20th-century American biologists 20th-century American women scientists 20th-century women mathematicians