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History of Flow Cytometry

History of Flow Cytometry

The first fluorescence-based flow cytometry device (ICP 11) was developed in 1968 by Wolfgang Göhde from the University of Münster, Germany and first commercialized in 1968/69 by German developer and manufacturer Partec through Phywe AG in Göttingen. At that time, absorption methods were still widely favored by other scientists over fluorescence methods [1]. The original name of the flow cytometry technology was pulse cytophotometry (German: Impulszytophotometrie). Only 10 years later in 1978, at the Conference of the American Engineering Foundation in Pensacola, Florida, the name was changed to flow cytometry, a term that quickly became popular. Soon after, flow cytometry instruments were developed, including the Cytofluorograph (1971) from Bio/Physics Systems Inc. (later: Ortho Diagnostics), the PAS 8000 (1973) from Partec, the first FACS instrument from Becton Dickinson (1974), the ICP 22 (1975) from Partec/Phywe and the Epics from Coulter (1977/78) [2].

Featured video

Introduction to Flow Cytometry

This introductory animation provides an overview of the four major systems of a flow cytometer and how they work together to analyze cell characteristics.


Historic Stains (1850-1953)

Up to 1850's
Only natural stains were available such as Saffron (which was what Leeuwenhoek used to stain muscle cells)

Used acidic and basic dyes to identify acidophilic, eosinophilic, basophilic and neutrophilic leukocytes 1880's to study the dynamics of ocular fluids- used fluorescein

August Köhler - 1904
Fluorescence UV Microscope

Pappenheim and Unna - early 1900's
Combined methyl green and pyronin to stain nuclei green and cytoplasm red

Robert Feulgen - 1925
Demonstrated that DNA was present in both animal and plant cell nuclei - developed a stoichiometric procedure for staining DNA involving a derivatizing dye, (fuchsin) to a Schiff base

Andrew Moldavan - 1934
Demonstrates use of a suspending fluid in which were blood cells - the measurements were made in a capillary tube using a photoelectric sensor to make extinction measurements
Manuscript: Photo-Electric Technique for the Counting of Microscopical Cells. Andrew Moldavan Montreal, Canada Science 80:188-189, 1934

Torbjorn Caspersson - (1938-1998) -Staining Nucleic acids
1941 - Demonstrated that "nucleic acids, far from being waste products, were necessary prerequisites for the protein synthesis in the cell (published in Naturwissenschaften in January 1941) and that they actively participated in those processes."
1950 - Demonstrated that both DNA and RNA increase in actively growing cells famous monograph in 1950 "Cell Growth and Cell Function" described nucleic acid and protein metabolism during normal and abnormal growth. - These studies were made using a Cadmium spark source for a UV light, and primitive electronic circuits for detection of signals.  Used Feulgen stain to stain nuclei.
Manuscript: Torbjorn O. Caspersson, History of the Development of Cytophotometry from 1935 to the present in Analytical and Quantitative Cytology and Histology, pp2-6, 1987.

Albert H. Coons, Hugh J. Creech and R. Norman Jones - 1941
Developed the fluorescence antibody technique - they labeled antipneumococcal antibodies with anthracene allowing them to detect both the organism and the antibody in tissue using UV excited blue fluorescence
Manuscript: Immunological Properties of an Antibody Containing a Fluorescent Group. Albert H. Coons, Hugh J. Creech and R. Norman Jones - Department of Bacteriology and Immunology, Harvard Medical School, and the Chemical Laboratory, Harvard University Proc. Soc. Exp.Biol.Med. 47:200-202, 1941.

Coons and Kaplan - 1950
Conjugated fluorescein with isocyanate - better blue green fluorescent signal - further away from tissue autofluorescence. This method used a very dangerous preparative step using phosgene gas

History of cellular clinical diagnostics
1943 Landmark monograph: Diagnosis of Uterine Cancer by the Vaginal Smear. Authored by George N. Papanicolaou (an anatomist) and Herbert F. Traut (a gynecologist).

Success of the Pap Smear
1941 26,000 deaths per year in the United States due to cancer of the uterus as reported by Papanicolaou and Trout.
1996 4,900 estimated deaths per year in the United States due to cervical cancer with nearly a 2-fold increase in population in the intervening half century. At least half of these deaths are women who never had a Pap smear.

1951 Cytoanalyzer by Airborne instruments of Mineola, New York. Utilized World War II technology. 1980s TICAS and CYBEST; Computer analysis and automated cytology projects.  (These pioneering systems proved insufficient for general use).

Oswald T. Avery - 1944
Demonstrated that DNA was the carrier of genetic information.

Gucker - 1947
History Of Flow Cytometry Gucker Photoelectric Counter for Colloidal Particles
Photoelectric Counter for Colloidal Particles

  • Developed a flow cytometer for detection of bacteria in aerosols
  • Published paper in 1947 (work was done during WWII and was classified).
  • Goal was rapid identification of airborne bacteria and spores used in biological warfare
  • Instrument: Sheath of filtered air flowing through a dark-field flow illuminated chamber. Light source was a Ford headlamp, PMT detector (very early use of PMT)

Early Cell Counter 1948
History Of Flow Cytometry Early Cell Counter
Early cell counter. Katherine Williams and C.S. Sanders (Atomic Energy Research Establishment) 1948 - Unclassified in 1956. and Trout.

Early Microfluorometric Scanner Robert Mellors 1951
History Of Flow Cytometry Early Microfluorometric Scanner Robert Mellors
R.C. Mellors & R. Silver, A microfluorometric scanner for the differential detection of cells: application to exfoliative cytology, Science 104, 1951

Two Color Cell Counter Patent 1953
History Of Flow Cytometry Two Color Cell Counter Patent
J.C. Parker and W.R. Horst 1953

P.J. Crosland-Taylor 1952-3
History Of Flow Cytometry P.J. Crosland-Taylor
Sheath Flow Principle - 1952-3
A Device for Counting Small Particles Suspended in a Fluid through a Tube P.J. Crosland-Taylor Bland-Sutton Institute of Pathology Middlesex Hospital, London, W.1. June 17, 1952 Nature 171: 37-38, 1953.

Watson & Crick-1953

History Of Flow Cytometry Watson & Crick
This is a picture of part of the original model built by Watson and Crick at Cambridge in 1953. The work which began with Avery's identification of DNA as the "transforming principle" thus led to research that overturned the old conception of DNA as a repetitive and simple molecule, confirmed DNA's role in genetic transmission, and, with James Watson and Francis Crick's 1953 paper, elucidated its structure. J. D. WATSON and F. H. C. CRICK A Structure for Deoxyribose Nucleic Acid Nature, 2 April 1953, VOL 171,737 1953

Blood Cell Counting - Pre-automation

  • The hemocytometer was the counting standard until the 1950's.
  • The dimension of this device was 3x3x0.1 mm. Typically RBCs were counted using a 1:200 dilution from the 1 x 106/mm3 in whole blood
  • Leukocytes (5x103/mm3) were diluted 1:10 in a lysing reagent and a dye to stain nuclei
  • Statistical variation is calculated by the following:
    • - The standard deviation of a count on n items in n1/2
    • - Considering no more than 500 cells could be possibly counted manually the standard deviation would therefore be 22
    • - The coefficient of Variation (CV) is 22/500 or 4.4%
    • - Add pipetting and dilution errors and it's about 10%

The Coulter Principle (1954-1955)

The Coulter Principle
As a particle passes through the aperture, it creates a resistance. The bigger the particle, the more the resistance, the greater the voltage. Each voltage spike is directly proportional to the size of the cell. Today every modern hematology analyzer depends in some way on the Coulter Principle.

History Of Flow Cytometry Wallace and Joseph Coulter
High Speed Automatic Blood Cell Counter and Cell Size Analyzer
History Of Flow Cytometry Coulter Counter

High Speed Automatic Blood Cell Counter and Cell Size Analyzer Wallace H. Coulter, Coulter Electronics, Chicago, Illinois. Proc.Natl.Electronics Conf.12:1034-1042, 1956


The First Coulter Counter

History Of Flow Cytometry The Coulter Principle
History Of Flow Cytometry Coulter Counter

The first commercial version of the Coulter Counter

History Of Flow Cytometry Patent  application

Coulter's Original 1953 Patent application

History Of Flow Cytometry Hand-drawn advertising drafts

Hand-drawn advertising drafts of the first Coulter Counter (1956)


Coulter Counter Model F

History Of Flow Cytometry Coulter Counter Model F

A method was devised for using the model F Coulter Counter for counting goat erythrocytes, which are smaller and more numerous than those of humans. Blood samples were taken from 25 goats, and the cells were counted using 100- and 70-micron aperature tubes. A visual count also was made of a portion of each sample. The results were analyzed statistically to determine which aperture would produce the most accurate and reproducible results when compared with the manual counts. It was found that counts obtained with the 100-micron aperature tube were not significantly different from the manual counts.

This technology found commercial success in the medical industry where it revolutionized the science of hematology. Red blood cells, white blood cells and platelets make up the majority of the formed elements in the blood. When whole anticoagulated human blood is diluted with isotonic saline, the Coulter principle can be applied to count and size the various cells that make up whole blood. The first commercial application of the Coulter principle to hematology came in 1954 with the release of the Coulter Counter Model A (developed by Wallace and brother Joseph R. Coulter.

Within a decade, literally every hospital laboratory in the United States had Coulter Counter, and today every modern hematology analyzer depends in some way on the Coulter Principle

Historical Overview (1956-1975)

von Bertalanffy & Bickis - 1956
The metachromatic fluorescence of AO was used to identify and quantitate RNA in tissues and that that normal and malignant cells could be discriminated.

Cytometry Analytic Techniques M.R. Mendelsohn - 1958
Pioneered photometry by the two-wavelength method with the appropriate mathematical manipulation of the transmittances.

Marylou Ingram - 1960's
Identified that radiation caused increase number of binucleated lymphocytes in peripheral blood - she used a scanner to detect these rare cells (1/10000).

Preston - 1964
Cytoanalyzer was designed to identify Ingram's rare cells using a Vidicon based system - digitized images of lymphocytes were produced stained with eosin-methylene azure dye combinations.

Early flow systems

Hallermann et al, Kosenow - 1964
AO staining of leukocytes - was able to use fluorescence (in a flow based system) to select leukocytes from red cells despite a low ratio (1/1000) because they took up lots of AO - also claimed to be able to discriminate between monocytes and PMN

History Of Flow Cytometry Molecular Formula
History Of Flow Cytometry Scan and structure

Lou Kamentsky - IBM -1963
LA Kamentsky & CN Liu, Computer-automated design of multifont print recognition logic, IBM J. Research & Development 7, 1963

History Of Flow Cytometry Lou Kamentsky
History Of Flow Cytometry Multifont print recognition

U.V. Scanning Measurements - 1963
Ultraviolet Absorption in Epidermoid Cancer Cells L.A. Kamentsky, H Derman and M. R. Melamed, Science 142, 1963

History Of Flow Cytometry Normal CellsHistory Of Flow Cytometry Normal Cells 2
History of Flow Cytometry-U.V. Scanning Measurements - 1963 (right)History Of Flow Cytometry Cancer Cells

Kamentsky System - 1963

History Of Flow Cytometry Dr. Kamentsky
History Of Flow Cytometry Dr. Melamed
History Of Flow Cytometry Dr. Koss

LA Kamentsky, MR Melamed & H. Derman, Spectrophotometer: New instrument for ultrarapid cell analysis, Science 150, 1965

1964 Kamentsky Sorter

History Of Flow Cytometry Kamentsky System

Diagram of the aparatus

1964 Kamentsky Sorter

History Of Flow Cytometry CD8+ Human T Cell

Spectrophotometric Cell Sorter; Louis A. Kamentsky1 and Myron R. Melamed2

  1. IBM Watson Laboratory, Columbia University, New York
  2. Memorial Sloan Kettering Cancer Center, New York

Science 9 June 1967: Vol. 156. no. 3780, pp. 1364 – 1365 ,DOI: 10.1126/science.156.3780.1364

1966 Kamentsky RCS: Four Sensors, Sorting, Auto Sampling and Computer Data Reduction

History Of Flow Cytometry 1966 Kamentsky RCS
History Of Flow Cytometry 1966 Kamentsky RCS 2

Two analytic instruments were built and one was delivered to LA Herzenberg at Stanford University 1967

Los Alamos Volume Sorter -1965

History Of Flow Cytometry Los Alamos Volume Sorter
History Of Flow Cytometry Los Alamos Volume Sorter 2
History Of Flow Cytometry Mack Fulwyler

Mack Fulwyler worked in Marvin van Dilla's lab at Los Alamos. He developed the sorter in 1965. He initially used electronic cell volume at Los Alamos National Labs. This instrument separated cells based on electronic cell volume (same principle as the Coulter counter) and used electrostatic deflection to sort. The cells sorted were RBC because they observed a bimodal distribution of cell volume when counting cells. The sorting principle was based on that developed for the inkjet printer by Richard Sweet at Stanford in 1965.

The mysterious red cell problem solved
So it was determined that RBC traveling through the orifice were identified as "different" only because of the rotation of the cells (which was essentially random) After determining that the bimodal distribution was artifactual, this group were able to sort neutrophils and lymphocytes from blood.

Los Alamos Flow Microfluorometer

History Of Flow Cytometry Los Alamos Flow Microfluorometer
History Of Flow Cytometry Marvin Van Dilla Working with Harry Crissman
History Of Flow Cytometry Fluorescent scatter
History Of Flow Cytometry Fluoreswcent spectra

Marvin Van Dilla Working with Harry Crissman at Los Alamos doing DNA analysis

Phywe AG of Gottingen - 1969
Produced the first commercial flow cytometer built around a Zeiss fluorescent microscope.

History Of Flow Cytometry ICP 11 (1969)

ICP 11 (1969) Distributed by Phywe, Göttingen The first commercial flow cytometer PDP 11 computer

History Of Flow Cytometry Wolfgang Göhde

Wolfgang Göhde


Lou Kamentsky - Biophysics Systems -1970
Bio/Physics Systems - 1970 commercial cytometer - the "Cytograph" He-Ne laser system at 633 nm for scatter (and extinction) - supposedly the first commercial instrument incorporating a laser. It could separate live and dead cells by uptake of Trypan blue. A fluorescence version called the "Cytofluorograph" followed using an air cooled argon laser at 488 nm excitation

Ortho Diagnostics (Johnson and Johnson) purchased Biophysics in 1976 and in 1977 the System 50 Cytofluorograph was developed - this was a droplet sorter, with a flat sided flow cell, forward and orthogonal scatter, extinction, 2 fluorescence parameters, multibeam excitation, computer analysis option. J&J exit business twice, mid 1980s and mid 1990s.

History Of Flow Cytometry Cytograph

ICP 11 (1969) Distributed by Phywe, Göttingen The first commercial flow cytometer PDP 11 computer

Herzenberg - Stanford - 1969
Len Herzenberg - Sorter based on fluorescence (arc lamp) built after working with one of Kamentsky's RCS systems where they built an instrument they called the Fluorescence Activated Cell Sorter (FACS)

History Of Flow Cytometry Len Herzenberg

ICP 11 (1969) Distributed by Phywe, Göttingen The first commercial flow cytometer PDP 11 computer

Herzenberg -1972 - Argon laser flow sorter - placed an argon laser onto their sorter and successfully did high speed sorting - Coined the term Fluorescence Activated Cell Sorting (FACS) This instrument could detect weak fluorescence with rhodamine and fluorescein tagged antibodies. A commercial version was distributed by B-D in 1974 and could collect forward scatter and fluorescence above 530 nm.

Particle Technology Inc. - COULTER -1971

  • Fulwyler began consulting for Coulter in the late 1960's. Spinning out LASL FCM and Particle manufacturing technologies.
  • In 1971, Mack Fulwyler resigned from LASL and established PTI as a Coulter subsidiary company
  • 1976 PTI dissolved, technology transferred to Florida
History Of Flow Cytometry Epics II 1975

Epics II 1975, Designed by Mack Fulwyler and Jim Corell Delivered to NCI/NIH

History Of Flow Cytometry TPS 1974 - 1979

TPS 1974 - 1979, Designed by Bob Auer


Hemalog D - 1974
Technicon - First commercial differential flow cytometer with light scatter and absorption at different wavelengths. Chromogenic enzyme substrates were used to identify neutrophils and eosinophils by peroxidase and monocytes by esterase, basophils were identified by the presence of glycosaminoglycans using Alcian Blue.  The excitation for all measurements was a tungsten-halogen lamp.

History Of Flow Cytometry Hemalog D

TPS 1974 - 1979, Designed by Bob Auer

Photo from Shapiro "Practical Flow Cytometry", 3rd. Ed.Wiley-Liss, 1994

Howard M. Shapiro - 1973-76
Shapiro and the Block instruments designed a series of multibeam flow cytometers that did differentials and multiple fluorescence excitation and emission

History Of Flow Cytometry Shapiro and the Block instruments

TPS 1974 - 1979, Designed by Bob Auer

LLNL High Speed Sorter - 1978
Marv Van Dilla and Phil Dean sorting chromosomes at LLNL around 1978, on the first fluorescence-based sorter developed there. The sorter shown was later modified to become the first dual-beam, fully computer-controlled, multi-parameter sorter. Father of the MoFlo.

History Of Flow Cytometry LLNL High Speed Sorter

TPS 1974 - 1979, Designed by Bob Auer


Invention of Monoclonal Antibody by Kohler and Milstein - 1975

History Of Flow Cytometry Kohler

1984 Nobel price for medicine: Niels K. Jerne, Georges J.F. Köhler, César Milstein "for theories concerning the specificity in development and control of the immune system and the discovery of the principle for production of monoclonal antibodies.

The referee reports were positive but cautious. The editors of Nature did not consider it of sufficient general interest to publish it as an article, and the original text had to be severely pruned to fit the length of a letter."Milstein commenting on his original paper in 1999

History Of Flow Cytometry Milstein

Interesting factoid

  • Len Herzenberg was doing a sabbatical in Cesar Milstein's laboratory in 1975 when Milstein had developed the Monoclonal Antibody.
  • It was Herzenberg that Milstein credited with coming up with a name for the cell than made the antibody - hybridoma!
  • The term hybridoma was proposed by Len Herzenberg during a sabbatical in my laboratory in 1976/1977. At a high-table conversation at a Cambridge College, Len was told by one of the dons that hybridoma was garbled Greek. By then however, the term was becoming popular among us, and we decided to stick to it." The hybridoma revolution: an offshoot of basic research Cesar Milstein, Bioessays, 21:966-973,1999.

Coulter Electronics (1976- 2009)

Epics V and 750 series - 1979 through 1985

A series of instruments which were essentially 5 watt argon ion laser instruments, complete with a multiparameter data analysis system, floppy drive and graphics printer



History Of Flow Cytometry EPICS V

Coulter's Original 1953 Patent application

EPICS V Dual Laser

History Of Flow Cytometry EPICS V Dual Laser

Hand-drawn advertising drafts of the first Coulter Counter (1956)



History Of Flow Cytometry EPICS 541

Coulter's Original 1953 Patent application


History Of Flow Cytometry EPICS 750

Hand-drawn advertising drafts of the first Coulter Counter (1956)


Stuart Schlossman

  • Schlossman at the Farber Institute in Boston, began to make monoclonal antibodies to white blood cell antigens in 1978. Eventually he collaborated with Ortho Diagnostics who distributed the famous "OK T4"etc., Mabs
  • Coulter Immunology also acquired rights to his antibodies

History of Coulter Hematology


1953 Model A

History Of Flow Cytometry 1953 Model A

Electronically measured cells

1968 Model S

History Of Flow Cytometry 1968 Model S

Completely Automated CBC


1977 S Plus

History Of Flow Cytometry 1977 S Plus

Completely Automated CBC

1985 VC

History Of Flow Cytometry 1985 VC

Integration of flow cytometry into a hematology analyzer


2000 Gen•S Cell

History Of Flow Cytometry 2000 Gen
  • AccuFlex
  • Decision Support Rules
  • IntelliKinetics
  • Automated Reticulocytes
  • Integrated SlideMaker and SlideStainer

1985 VC

History Of Flow Cytometry Hematology LH 700 Series
  • Random Access
  • AccuCount Technology
  • Extended Linearity
  • Decreased need for Manual Intervention
  • WBC Interference Correction
  • NRBC enumeration

History of Coulter Flow Cytometry


1975 TPS

History Of Flow Cytometry 1975 TPS

Two Parameter Cell Sorting

1984 Epics C

History Of Flow Cytometry 1984 Epics C

Clinical Flow Sorter


1986 Epics PROFILE

History Of Flow Cytometry 1986 Epics PROFILE

Clinical Flow Analyzer

1987 Q Prep

History Of Flow Cytometry Automated Sample Prep

Automated Sample Prep


1988 Cyto-Stat Monoclonal Antibodies

History Of Flow Cytometry 1988 Cyto-Stat
  • Ready to use
  • No Wash
  • Single and Multicolor Antibodies


History Of Flow Cytometry 1993 EPICS XL / XL-MCL
  • Bench-top analyzer
  • Four color analysis
  • Autoloader for samples
  • Digital Signal Processing

Flow Cytometers

Modern flow cytometers are able to analyze several thousand particles every second, in "real time," and can actively separate and isolate particles having specified properties. A flow cytometer is similar to a microscope, except that, instead of producing an image of the cell, flow cytometry offers "high-throughput" (for a large number of cells) automated quantification of set parameters. To analyze solid tissues, single-cell suspension must first be prepared.

Early flow cytometers were, in general, experimental devices, but recent technological advances have created a considerable market for the instrumentation, as well as the reagents used in analysis, such as fluorescently-labeled antibodies and analysis software. Modern instruments usually have multiple lasers and fluorescence detectors (the current record for a commercial instrument is 4 lasers and 18 fluorescence detectors).

History Of Flow Cytometry Analysis of a marine sample History Of Flow Cytometry Analysis of a marine sample 2 History Of Flow Cytometry Analysis of a marine sample 3

Analysis of a marine sample of photosynthetic picoplankton by flow cytometry showing three different populations (Prochlorococcus, Synechococcus, and picoeukaryotes)

Increasing the number of lasers and detectors allows for multiple antibody labeling, and can more precisely identify a target population by their phenotype. Certain instruments can even take digital images of individual cells, allowing for the analysis of fluorescent signal location within or on the surface of cells. The data generated by flow-cytometers can be plotted in a single dimension, to produce a histogram, or in two-dimensional dot plots or even in three dimensions. The regions on these plots can be sequentially separated, based on fluorescence intensity, by creating a series of subset extractions, termed "gates." Specific gating protocols exist for diagnostic and clinical purposes especially in relation to hematology. The plots are often made on logarithmic scales. Because different fluorescent dyes' emission spectra overlap [1], signals at the detectors have to be compensated electronically as well as computationally. Often, data accumulated using the flow cytometer can be re-analysed (using software, e.g., Kaluza™ [5]) elsewhere, freeing up the machine for other people to use.

Commercial Instruments
Beckman Coulter's complete range of automation and information systems help you streamline processes for maximum efficiency. From delivery of more timely, accurate and reliable patient test results to the elimination of bottlenecks, our automation and information system solutions empower you to manage lab operations more efficiently and cost-effectively.

Flow Cytometry - Definitions


Flow Cytometry (cyto=cell) (metry=measurement)
Measuring properties of cells in a flowing system

Flow Sorting
Sorting (physically separating) cells based on properties measured in a flowing system

Flow Cytometer
Early flow cytometers were, in general, experimental devices, but recent technological advances have created a considerable market for the instrumentation, as well as the reagents used in analysis, such as fluorescently-labeled antibodies and analysis software. Modern instruments usually have multiple lasers and fluorescence detectors.

Principle of flow cytometry


A beam of light (usually laser light) of a single wavelength is directed onto a hydrodynamically-focused stream of fluid. A number of detectors are aimed at the point where the stream passes through the light beam: one in line with the light beam (Forward Scatter or FSC) and several perpendicular to it (Side Scatter (SSC) and one or more fluorescent detectors).

Each suspended particle from 0.2 to 150 micrometers passing through the beam scatters the light in some way, and fluorescent chemicals found in the particle or attached to the particle may be excited into emitting light at a longer wavelength than the light source. This combination of scattered and fluorescent light is picked up by the detectors, and, by analyzing fluctuations in brightness at each detector (one for each fluorescent emission peak), it is then possible to derive various types of information about the physical and chemical structure of each individual particle.


  1. "Wallace H. Coulter 1913-1998". Beckman Coulter, Inc. Retrieved on 2008-07-31.
  2. "Joseph R. Coulter Jr. 1924-1995". Beckman Coulter, Inc. Retrieved on 2008-07-31.
  3. Presentation by Bob Auer - 2 December 2008 - Images and text.
  4. Kamentsky, Proceedings of the Conference "Cytology Automation" in Edinburgh, 1970
  5. Images of non Beckman Coulter equipment are from J. Paul Robinson lectures posted on the Purdue website.
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