The history

The history of understanding taste

From Aristotle and the six rasas of Ayurveda to a myth-making tongue map, the discovery of umami, and receptors for every taste.

People have argued about how many tastes there are for over 2,000 years. For most of that time taste was a matter of philosophy and cooking. Microscopes found the taste buds, careful tests found (and a misreading invented) a tongue map, and a Japanese chemist found a taste Europe had missed. In the last 30 years, genetics revealed the receptor for each of the five tastes, and showed that chilli heat is really a pain and heat sensor at work.

2,300+
years
26
moments
10
people
8
places

c. 350 BCE

Written list of basic tastes

Aristotle, Greece

c. 2nd century BCE to 2nd century CE

Six tastes as the basis of a healthy diet

Charaka Samhita, India

1867

Taste buds seen under the microscope

Christian Lovén and Gustav Schwalbe

1908

Umami identified

Kikunae Ikeda, Japan

1912

Chilli heat scale

Wilbur Scoville, USA

1997

Heat and capsaicin sensor (TRPV1) found

David Julius, USA

2007

Chilli certified above one million SHU

Bhut jolokia, north-east India

2018–2019

Sour receptor (OTOP1) identified

Liman and Zuker labs, USA

c. 350 BCENaming the tastes

400 BCE – 1800

Naming the tastes

Philosophers and physicians in Greece and India sort tastes into short lists, including some, like pungent and astringent, that science now calls touch.

350 BCE

c. 350 BCE

Aristotle lists the tastes

AristotleAthens, Greece

In his book On the Soul, Aristotle wrote that the two basic tastes are sweet and bitter. Between them he placed oily, salty, pungent, harsh (astringent) and sour, a list of seven or eight flavours.

Why it mattered. It was one of the first attempts to sort tastes into a short list of basics, an idea still used today.

100 BCE

c. 2nd century BCE to 2nd century CE

Ayurveda names six rasas

The Charaka Samhita (compiled by physicians in the tradition of Agnivesha and Charaka)Ancient India

The Charaka Samhita, a founding text of Ayurveda, describes six rasas, or tastes: madhura (sweet), amla (sour), lavana (salty), katu (pungent), tikta (bitter) and kashaya (astringent). A healthy diet, it says, balances all six. Its date is uncertain; it was revised over several centuries.

Why it mattered. Its six tastes include pungent and astringent, which science now files under touch and pain rather than taste, as this box shows.

1860 – 1975

Buds, maps and myths

Microscopes reveal the taste buds; a study of the tongue edge is misread into the famous tongue map, then tested and found wrong.

1867

Taste buds are discovered

Christian Lovén and Gustav SchwalbeStockholm, Sweden, and Germany

Working separately, the Swedish anatomist Christian Lovén and the German anatomist Gustav Schwalbe looked at the tongue under the microscope and found small onion-shaped clusters of cells, which they called taste bulbs or taste goblets.

Why it mattered. It showed that taste is sensed by special groups of cells, not by the whole tongue surface.

1901

Hänig measures the edge of the tongue

David P. HänigLeipzig, Germany

In a paper called "On the psychophysics of the sense of taste", Hänig dripped sweet, sour, salty and bitter solutions on spots around the edge of people's tongues and found the weakest strength each could notice. The differences between spots were small.

Why it mattered. His careful data were later misread as a tongue map, one of the most famous myths in science teaching.

1942

A graph without units starts a myth

Edwin G. BoringHarvard University, USA

In his history of the psychology of the senses, Boring redrew Hänig's 1901 results as a graph of how sensitive each part of the tongue edge was, with no numbers on the scale. Readers took the peaks to mean each taste lived in only one area, and a tongue map spread into textbooks.

Why it mattered. It shows how a small misreading can travel for decades.

1965

Taste cells are always being replaced

Lloyd Beidler and Richard SmallmanFlorida State University, USA

By labelling new cells in rats, Beidler and Smallman showed that taste-bud cells are constantly replaced, living about ten days on average before new ones take their place.

Why it mattered. It explains why taste usually recovers after a burn from hot food or drink.

1974

The tongue map is tested and fails

Virginia B. CollingsUniversity of Pittsburgh, USA

Collings repeated the tests on many spots of the tongue and found that all four tastes then known could be sensed everywhere with taste buds. The weakest noticeable strength differed a little from place to place, but no area was blind to any taste.

Why it mattered. It was the scientific end of the tongue map, even if school books kept it for years.

1900 – 1995

Umami, chillies and genes

A fifth taste is found in seaweed broth, chilli heat gets a scale, and some people turn out to be blind to a bitter chemical.

1908

Ikeda finds umami in kombu

Kikunae IkedaTokyo Imperial University, Japan

The chemist Kikunae Ikeda wondered why kombu seaweed broth (dashi) tasted so savoury. He boiled down large amounts of kombu and extracted glutamic acid, whose salts gave the taste. He named it umami, from the Japanese for "delicious".

Why it mattered. It was the first scientific claim for a fifth basic taste, though the West took most of a century to accept it.

1909

MSG goes on sale

Kikunae Ikeda and Saburosuke SuzukiJapan

Ikeda patented a way to make monosodium glutamate (MSG) and teamed up with the businessman Saburosuke Suzuki to sell it as a seasoning called Ajinomoto, "essence of taste".

Why it mattered. It turned a taste discovery into one of the world's most widely used food ingredients.

1912

Scoville measures chilli heat

Wilbur ScovilleDetroit, USA

The pharmacist Wilbur Scoville, working for the drug company Parke-Davis, soaked dried chillies in alcohol, then diluted the extract with sugar water until a panel of tasters could no longer feel the heat. The number of dilutions became the Scoville heat unit.

Why it mattered. It gave the world a scale for chilli heat, still used today, though labs now measure capsaicin with machines.

1931

The chemical some people cannot taste

Arthur FoxDuPont laboratory, Wilmington, USA

Fox was pouring a powder called PTC when some of it blew into the air. His colleague C. R. Noller complained it was very bitter; Fox, standing closer, tasted nothing. Testing others showed that people differ: roughly a third of people worldwide cannot taste it.

Why it mattered. It was the first clear sign that taste is partly in our genes. The gene, TAS2R38, a bitter receptor, was found in 2003.

1985

Umami becomes a scientific word

First Umami International SymposiumHawaii, USA

Researchers from many countries met in Hawaii for the first international meeting on umami. Shizuko Yamaguchi showed with careful taste tests that umami could not be made by mixing the other four tastes, and "umami" was adopted as the scientific name for the taste of glutamate.

Why it mattered. It put umami on the world's scientific map, though its receptor was still unknown.

1994

Supertasters

Linda Bartoshuk and colleaguesYale University, USA

Bartoshuk's team found that people fall into three groups for the bitter chemical PROP: non-tasters, medium tasters and "supertasters", who find it intensely bitter and tend to have more fungiform papillae. In their samples, about a quarter of people were supertasters.

Why it mattered. It showed that people really do live in different taste worlds.

1997 – 2019

A receptor for every taste

Genetics finds the sensors: T1Rs for sweet and umami, T2Rs for bitter, ENaC for salt, OTOP1 for sour, and TRP channels for heat and cold.

1997

The capsaicin receptor is found

David Julius and colleaguesUniversity of California, San Francisco, USA

Julius's team searched through thousands of genes for one that made cells respond to capsaicin. They found TRPV1, a channel that also opens at painful heat above about 43 °C.

Why it mattered. It explained why chilli feels hot, and opened the science of how we sense temperature.

1999

The first taste receptor genes

Mark Hoon, Nicholas Ryba, Charles Zuker and colleaguesNIH and University of California, San Diego, USA

The team found two genes for receptors made only in taste cells, later named T1R1 and T1R2. It took two more years to learn what they detect.

Why it mattered. It was the first step to naming the molecules behind each taste.

2000

A family of bitter receptors

Several teams, including Zuker and Ryba, and Linda BuckUSA

Three papers described the T2Rs, a family of about 25 bitter receptors in humans. Many bitter cells carry several kinds, so one cell can warn of many different poisons. The same year, a team in Miami proposed a glutamate receptor for umami.

Why it mattered. It showed why bitter is so sensitive: it is a warning system built to catch thousands of chemicals.

2001

2001–2002

Sweet and umami receptors are pinned down

Greg Nelson, Charles Zuker, Nicholas Ryba and othersUSA

Teams found a third gene, T1R3, and showed that T1R2 + T1R3 together make the sweet receptor (2001), while T1R1 + T1R3 make the umami receptor for glutamate (2002).

Why it mattered. Umami finally had a receptor of its own, and its place as the fifth basic taste was settled.

2002

The cold and menthol sensor

David Julius's and Ardem Patapoutian's labsSan Francisco and La Jolla, USA

Two teams independently found TRPM8, a channel on nerve endings that opens with cool temperatures and with menthol from mint.

Why it mattered. It explained why mint feels cold even when it is not.

2010

The salt channel

Jayaram Chandrashekar, Charles Zuker, Nicholas Ryba and colleaguesUSA

Mice without the sodium channel ENaC in their taste cells lost their liking for low salt. So ENaC is the sensor for pleasant saltiness, while very salty food also triggers bitter and sour cells.

Why it mattered. It explained why a little salt tastes good and a lot tastes bad.

2015

Is fat a sixth taste?

Cordelia Running, Bruce Craig and Richard MattesPurdue University, USA

Taste tests suggested that fatty acids have a taste of their own, which the team proposed calling "oleogustus". Earlier work had found fat-sensing receptors such as CD36 on taste cells. Scientists still debate it, along with "kokumi", a richness linked to a calcium-sensing receptor.

Why it mattered. The list of basic tastes may not be finished.

2018

2018–2019

The sour receptor at last

Emily Liman's lab; Charles Zuker's labLos Angeles and New York, USA

Sour was the last basic taste without a known receptor. In 2018 Liman's team showed that OTOP1, a channel that lets protons in, is found in sour cells. In 2019 her lab and Zuker's showed that mice without it barely sense sour.

Why it mattered. It completed the set: each of the five tastes now has a known sensor.

2007 – today

Records, a pandemic and a Nobel

Chilli breeders chase heat records, COVID-19 shows the world how flavour depends on smell, and the heat sensor wins a Nobel Prize.

2007

February 2007

The first million-Scoville chilli

Paul Bosland, Chile Pepper InstituteNew Mexico State University, USA, with seeds from north-east India

Guinness World Records certified the bhut jolokia, the "ghost pepper" grown in Assam and nearby states, as the world's hottest chilli, after lab tests measured 1,001,304 Scoville heat units.

Why it mattered. It was the first chilli certified above a million SHU, nearly double the previous record holder.

2013

November 2013

The Carolina Reaper

Ed Currie, PuckerButt Pepper CompanySouth Carolina, USA

Guinness World Records confirmed the Carolina Reaper, a cross-bred chilli, as the hottest, at an average of 1,569,300 Scoville heat units; a 2017 re-test put it at 1,641,183.

Why it mattered. Breeders were now deliberately crossing chillies to push the heat higher.

2020

Losing taste and smell to COVID-19

Doctors and health agencies worldwideWorldwide

During the COVID-19 pandemic, sudden loss of smell, and with it much of the sense of taste, became one of the best-known symptoms. Health services such as the UK's NHS and the US CDC added it to their symptom lists in 2020.

Why it mattered. Millions of people learned how much of flavour depends on smell, and research into smell loss grew fast.

2021

October 2021

A Nobel Prize for heat and touch

David Julius and Ardem PatapoutianStockholm, Sweden

The Nobel Prize in Physiology or Medicine went to Julius, for finding TRPV1 with capsaicin, and to Patapoutian, for finding the touch sensors PIEZO1 and PIEZO2. Both also found TRPM8, the cold sensor.

Why it mattered. The science behind chilli heat and minty cool reached the top of medicine.

2023

Pepper X

Ed Currie, PuckerButt Pepper CompanySouth Carolina, USA

Guinness World Records named Pepper X, bred by the grower of the Carolina Reaper over a decade, the hottest chilli, at an average of 2,693,000 Scoville heat units.

Why it mattered. Chilli breeding keeps pushing a plant's defence chemical to extremes.

By the numbers

The hottest chilli on record

Guinness-certified record holders: from the first million-SHU chilli in 2007, breeders have pushed the heat up by about two and a half times.

1000000 Scoville heat units1500000 Scoville heat units2000000 Scoville heat units2500000 Scoville heat units3000000 Scoville heat units 201020152020 2007: Bhut jolokia (India): 1,001,304 SHU20072011: Trinidad Scorpion Butch T (Australia): 1,463,700 SHU, after the Naga Viper briefly held 1,382,11820112013: Carolina Reaper (USA): 1,569,300 SHU20132017: Carolina Reaper re-tested: 1,641,183 SHU20172023: Pepper X (USA): 2,693,000 SHU2023
  1. 2007 Bhut jolokia (India): 1,001,304 SHU
  2. 2011 Trinidad Scorpion Butch T (Australia): 1,463,700 SHU, after the Naga Viper briefly held 1,382,118
  3. 2013 Carolina Reaper (USA): 1,569,300 SHU
  4. 2017 Carolina Reaper re-tested: 1,641,183 SHU
  5. 2023 Pepper X (USA): 2,693,000 SHU

Did you know?

The tongue map, with sweet on the tip and bitter at the back, is a myth: every part with taste buds tastes all five.

You can notice bitter quinine at a strength over 1,000 times weaker than the weakest sugar you can notice.

Chilli heat is not a taste: capsaicin opens TRPV1, the same sensor that warns of burning heat above about 43 °C.

Your taste cells are replaced about every ten days, some after a few weeks.

Ayurveda counted six tastes about 2,000 years ago; two of them, pungent and astringent, are now known to be touch and pain rather than taste.

The people

Who figured it out

Aristotle

384 – 322 BCE · Philosopher · Greece

Made one of the first lists of basic tastes, with sweet and bitter as the extremes.

Kikunae Ikeda

1864 – 1936 · Chemist · Japan

Found glutamate in kombu broth and named its taste umami.

Wilbur Scoville

1865 – 1942 · Pharmacist · USA

Invented the dilution test that still gives chilli heat its units.

Arthur L. Fox

active 1930s · Chemist · USA

Discovered by accident that some people cannot taste the bitter chemical PTC.

Virginia B. Collings

20th century · Psychologist · USA

Tested the tongue map in 1974 and showed that every area with taste buds senses every taste.

Linda Bartoshuk

born 1938 · Taste scientist · USA

Found supertasters and showed how differently people experience taste.

Charles Zuker and Nicholas Ryba

Zuker born 1957 · Neuroscientists · Chile and the UK, working in the USA

Their labs found or pinned down receptors for sweet, umami, bitter, salt and sour.

Emily Liman

20th–21st century · Neuroscientist · USA

Her lab identified OTOP1, the proton channel behind sour taste.

David Julius

born 1955 · Physiologist · USA

Used capsaicin to find TRPV1, the heat sensor; Nobel Prize 2021.

Ardem Patapoutian

born 1967 · Neuroscientist · Lebanon, working in the USA

Found the touch sensors PIEZO1 and PIEZO2 and co-discovered the cold sensor TRPM8; Nobel Prize 2021.

Where it happened

8 places, one idea

Sources

Where this comes from

Dates marked “c.” are approximate, and historians sometimes disagree about who was first. If you spot a mistake, tell us.

  1. Taste Wikipedia
  2. On the Soul (De Anima), Book II Internet Classics Archive, MIT
  3. Rasa Vimana Charak Samhita Online (NIIMH / CCRAS)
  4. Charaka Samhita Wikipedia
  5. Physiology of the senses: a prominent area of science in Uppsala at the end of the nineteenth century Upsala Journal of Medical Sciences (PMC)
  6. Taste (1911 Encyclopaedia Britannica) Wikisource
  7. Tongue map Wikipedia
  8. The taste map of the tongue you learned in school is all wrong Smithsonian Magazine
  9. Umami Wikipedia
  10. Umami: Innovation story Ajinomoto Group
  11. Monosodium glutamate Wikipedia
  12. Scoville scale Wikipedia
  13. Wilbur Scoville Wikipedia
  14. Phenylthiocarbamide Wikipedia
  15. The relationship between chemical constitution and taste (PNAS 18:115, 1932) PNAS
  16. Renewal of cells within taste buds (J Cell Biol 27:263, 1965) Journal of Cell Biology
  17. Taste bud Wikipedia
  18. Human taste response as a function of locus of stimulation on the tongue and soft palate (Percept Psychophys 16:169) Perception & Psychophysics (Springer)
  19. What is umami? Umami Information Center
  20. Supertaster Wikipedia
  21. PTC/PROP tasting: anatomy, psychophysics, and sex effects (Physiol Behav 56:1165) Physiology & Behavior
  22. The capsaicin receptor: a heat-activated ion channel in the pain pathway (Nature 389:816) Nature
  23. Press release: The Nobel Prize in Physiology or Medicine 2021 NobelPrize.org
  24. Putative mammalian taste receptors (Cell 96:541, 1999) Cell
  25. The receptors and cells for mammalian taste (Nature 444:288, 2006) Nature
  26. A novel family of mammalian taste receptors (Cell 100:693, 2000) Cell
  27. Taste receptor Wikipedia
  28. Mammalian sweet taste receptors (Cell 106:381, 2001) Cell
  29. An amino-acid taste receptor (Nature 416:199, 2002) Nature
  30. Identification of a cold receptor reveals a general role for TRP channels in thermosensation (Nature 416:52) Nature
  31. TRPM8 Wikipedia
  32. Bhut jolokia Wikipedia
  33. 'Bhut Jolokia': the world's hottest known chile pepper (HortScience 42:222) HortScience
  34. The cells and peripheral representation of sodium taste in mice (Nature 464:297) Nature
  35. Epithelial sodium channel Wikipedia
  36. Confirmed: Smokin Ed's Carolina Reaper sets new record for hottest chilli Guinness World Records
  37. Carolina Reaper Wikipedia
  38. Oleogustus: the unique taste of fat (Chem Senses 40:507) Chemical Senses
  39. An evolutionarily conserved gene family encodes proton-selective ion channels (Science 359:1047) Science
  40. Sour sensing from the tongue to the brain (Cell 179:392) Cell
  41. Symptoms of COVID-19 NHS
  42. Taste disorders NIDCD, US National Institutes of Health
  43. David Julius Wikipedia
  44. Hottest chili pepper Wikipedia
  45. Pepper X dethrones Carolina Reaper as world's hottest chilli pepper Guinness World Records
  46. Pepper X Wikipedia

That's the history. Now see how it works.