日本語版

tomato

ingredient / ingredient-tomato

トマト / Solanum lycopersicum

Tomato is the edible tomato fruit. Ripe fruit of some wild relatives is bitter because bitter steroidal glycoalkaloids, including α-tomatine, remain.

Photographs (captions in Japanese)

6 small oval tomatoes in 2 rows on a green stem, showing a color gradient from red to orange to green from the top. The background is blurred green leaves.

Cherry tomatoes in one cluster with fruit at different ripening stages, green, orange, and red.

Shown: Solanum lycopersicumPart / state: Fruit (in clusters, including ripening stages from green to red)

Photo: Juan Carlos Fonseca Mata / CC BY-SA 4.0 / original image / ledger's original image

Scope and provenance of the photo

Identification: Identified as labelled in the source. The description is "Maduración del tomate (Solanum lycopersicum)" (ripening of the tomato). I opened the image and confirmed that red, orange, and green fruits are lined up in stages in the same cluster.

Image shown: The original from Wikimedia Commons, reduced to 2400 px on the long side. No cropping or color changes. For display, a WebP capped at 1200 px wide is served; the ledger's original image can be checked from the link above.

On this page (8 sections)
  1. Overview
  2. Sources for this overview (2)
  3. Topic hubs
  4. Other names
  5. Follow the connections
  6. Related entries (5)
  7. Composition and content (1)
  8. Mechanism and receptors (3)
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Reading the bitterness

Fully ripe tomato fruit is generally not bitter, owing in part to a chemical change mediated by a specific enzyme. GAME31, a 2-oxoglutarate-dependent dioxygenase (2-ODD), hydroxylates the bitter steroidal glycoalkaloid α-tomatine to hydroxytomatine. Starting with this reaction, a pathway during ripening converts it into nonbitter compounds such as esculeoside A, removing bitterness from ripe fruit, according to LC-MS metabolite analysis and experiments manipulating gene expression. Because this change occurs during ripening near harvest, it provides a harvest-stage example relevant to how bitterness changes with cultivation and harvest timing.

The pathway’s effect on composition was also tested in reverse. In a line in which GAME31 activity was suppressed by silencing, steroidal glycoalkaloids that normally remain only in trace amounts in ripe fruit increased by as much as 22-fold relative to wild type. No human sensory test was performed: attributing bitterness to these compounds rests on their known bitter properties, not on measured taste in these fruit. Chemical analysis did confirm that altering this one enzyme, GAME31, substantially changes ripe-fruit composition.

A separate enzyme, Sl23DOX, a type of 2-oxoglutarate-dependent dioxygenase, has also been identified as hydroxylating α-tomatine at position 23. Within the scope of the abstract, biochemical analysis of recombinant Sl23DOX protein and confirmation in transgenic tomato plants overexpressing or silencing this enzyme show that it catalyzes 23-hydroxylation; the product spontaneously isomerizes to neorickiioside B, an intermediate of ripening-stage metabolism. How this pathway coordinates with or substitutes for the GAME31 pathway is not clear from the abstract.

According to the abstract, α-tomatine accumulates in all tomato tissues, particularly at high concentrations in leaves and immature green fruit, and is described as a compound that causes bitterness. This is background in a recombinant-protein study, however, without specific concentration values or human sensory evaluation.

Sources for this overview

  1. Pathways to defense metabolites and evading fruit bitterness in genus Solanum evolved through 2-oxoglutarate-dependent dioxygenases

    Cárdenas PD; Sonawane PD; Heinig U; ほか16名 / 2019 / journal article / full text checked

    Checked for this overview: Used the description of the GAME31 hydroxylation pathway. / Used the result while explicitly noting the absence of sensory testing.

  2. Identification of α-Tomatine 23-Hydroxylase Involved in the Detoxification of a Bitter Glycoalkaloid

    Nakayasu M; Akiyama R; Kobayashi M; Lee HJ; Kawasaki T; Watanabe B; Urakawa S; Kato J; Sugimoto Y; Iijima Y; Saito K; Muranaka T; Umemoto N; Mizutani M / 2020 / journal article / abstract checked

    Checked for this overview: The position-23 hydroxylation pathway through Sl23DOX, distinct from GAME31. / Tissue distribution of α-tomatine, particularly high in leaves and immature green fruit.

Other names: Solanum lycopersicum (la)

Follow the connections

tomato →contains→ α-tomatine →— sensory attribute→ bitterness

α-tomatine →occurs in→ tomato

Structured claims — open a section to read each claim's confidence, evidence and limits

Composition and content (1)

tomato contains α-tomatinepreliminary

Scope: Layer: food. The background description of α-tomatine tissue distribution and taste in a recombinant-protein study.

Note: Only the abstract was checked; the full text was not reached. The abstract gives no specific concentration values. Earlier claims concern the GAME31 mechanism by which ripe fruit loses bitterness; this claim adds the association of high concentration and bitterness in different tissues, immature fruit and leaves.

Summarising earlier studies, it states that α-tomatine accumulates in all tomato tissues, especially leaves and immature green fruit, and causes a bitter taste.
Limitations: Background summary, not a main result of this study; the study has no human sensory evaluation.
Source: Identification of α-Tomatine 23-Hydroxylase Involved in the Detoxification of a Bitter Glycoalkaloid (Nakayasu M; Akiyama R; Kobayashi M; Lee HJ; Kawasaki T; Watanabe B; Urakawa S; Kato J; Sugimoto Y; Iijima Y; Saito K; Muranaka T; Umemoto N; Mizutani M, 2020) · abstract checked
Show dataingredient-tomato —contains→ compound-alpha-tomatine
claim: clm-w71b1-tomato-alphatomatine-tissue-distribution / type: composition / updated: 2026-09-07

Mechanism and receptors (3)

tomato has debittering most likely due to Hydroxylation of the bitter steroidal glycoalkaloid alpha-tomatine to hydroxytomatine by GAME31 (a 2-oxoglutarate-dependent dioxygenase). During ripening, a pathway starting from this reaction converts it into non-bitter esculeoside A and related compounds, which removes the bitterness of the ripe fruit.supported

Scope: Tomato, Solanum lycopersicum (cultivated varieties including cv. M82)

Note: In lines where GAME31 was silenced, SGA in ripe fruit increased up to 22-fold relative to the wild type, confirming through reverse genetics that loss of this enzyme can bring bitterness back.

Showed, using LC-MS metabolite profiling and gene-expression manipulation lines, that the 2-ODD enzyme GAME31 hydroxylates alpha-tomatine to hydroxytomatine, forming the first step of the pathway that converts it into non-bitter esculeoside A and related compounds during ripening.
Limitations: No human sensory testing was performed. The attribution of bitterness is based on chemical structure (that these are known bitter compounds).
In lines where GAME31 was silenced, SGA -- normally present only in trace amounts in ripe fruit -- increased up to 22-fold relative to the wild type. Confirmed through reverse genetics that loss of GAME31 function leads to SGA accumulation (bitterness).
Limitations: No sensory evaluation was performed. Chemical quantification only.
Show dataingredient-tomato —has_debittering_most_likely_due_to→ (the claim sentence above)
claim: clm-tomato-game31-debitters-ripe-fruit / type: mechanism / updated: 2026-09-02
tomato is processed by Sl23DOX, an α-tomatine 23-hydroxylase and a 2-oxoglutarate-dependent dioxygenase, hydroxylates position 23. Its product spontaneously isomerizes to neorickiioside B, forming an intermediate in α-tomatine metabolism during ripening. This is a parallel metabolic pathway involving an enzyme distinct from GAME31.preliminary

Scope: Layer: food. Biochemical analysis with recombinant Sl23DOX protein and confirmation in Sl23DOX-overexpressing and silenced transgenic tomato plants.

Recombinant Sl23DOX protein hydroxylated α-tomatine at position 23, and the product spontaneously isomerised to neorickiioside B, a metabolic intermediate that appears during ripening; metabolites were analysed in overexpressing and silenced tomato plants.
Limitations: Only the abstract was checked. No human sensory test was performed; attribution of bitterness is based on compound identification.
Source: Identification of α-Tomatine 23-Hydroxylase Involved in the Detoxification of a Bitter Glycoalkaloid (Nakayasu M; Akiyama R; Kobayashi M; Lee HJ; Kawasaki T; Watanabe B; Urakawa S; Kato J; Sugimoto Y; Iijima Y; Saito K; Muranaka T; Umemoto N; Mizutani M, 2020) · abstract checked
Show dataingredient-tomato —is_processed_by→ (the claim sentence above)
claim: clm-w71b1-tomato-sl23dox-pathway / type: mechanism / updated: 2026-09-07
α-tomatine occurs in tomatosupported

Scope: Layer: food. Comparison of red-ripe fruit from NOR-KO and RIN-KO tomato mutants with wild type; glycoside quantification by LC-MS and GAME gene-expression analysis by RT-qPCR.

In NOR-KO and RIN-KO fruits, compared with WT fruits, the levels of α-tomatine, hydroxytomatine, acetoxytomatine, and acetoxy-hydroxytomatine were significantly increased, while esculeoside A was markedly lower.
Limitations: These data come from gene-knockout mutants under manipulated conditions beyond natural variation in ordinary ripening. The sensory intensity of α-tomatine bitterness itself was not measured in this study; it is treated as a known property based on earlier literature.
Source: Removal of toxic steroidal glycoalkaloids and bitterness in tomato is controlled by a complex epigenetic and genetic network (Bai F; Wu M; Huang W; Xu W; Wang Y; Zhang Y; Zhong Z; Hong Y; Pirrello J; Bouzayen M; Liu M, 2025) · full text checked
Show datacompound-alpha-tomatine —occurs_in→ ingredient-tomato
claim: clm-w72a1-tomato-nor-rin-alphatomatine-retention / type: mechanism / updated: 2026-09-08

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