Paper · 2025 · Frontiers in Plant Science

Alternate wetting and drying irrigation at tillering stage enhances the heat tolerance of rice by increasing sucrose and cytokinin content in panicles

Ti Gao; Keran Xie; Qiuqian Hu; et al.

17
claims
51
verbatim quotes
8
citations
4
research areas
✓
Open access · cc-by

Abstract

High temperature events have occurred frequently in recent years under global warming conditions. High temperature during panicle initiation (PI) poses significant negative impacts on rice grain yield. Alternate wetting and drying irrigation (AWD) is widely adopted in rice cultivation. Here, to investigate the alleviating effect of AWD on heat damage to rice ( Oryza sativa L.), we selected four rice varieties with different high-temperature resistance (Liangyoupeijiu, IR64, Huanghuazhan, Shanyou 63), two irrigation treatments at the tillering stage including AWD and continuous flooding (CF) and two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK) were applied. HDT significantly reduced the yield of all four varieties except for Shanyou 63, primarily by decreasing the spikelet fertility and spikelet number, while AWD significantly improved the two traits under HDT. Moreover, HDT reduced the photosynthetic rate, increased the starch content in leaves and stems, and decreased the transport of sucrose to panicles. AWD reduced the panicle temperature, promoted sucrose synthesis in leaves, increased sucrose content and sucrose hydrolase activity in panicles under HDT. HDT also increased the content of abscisic acid and decreased that of cytokinins (CTKs), indole-3-acetic acid (IAA), and gibberellins (GAs) in panicles under CF. AWD increased the expression of CTK biosynthesis genes ( IPT , CYP735A , LOG ) and decreased that of CTK degradation gene CKX in roots and panicles under HDT, while enhanced the xylem sap flow rate and contents of CTKs, IAA, and GAs in panicles. Furthermore, AWD exhibited a more pronounced alleviating effect on HDT damage in heat-sensitive varieties than in heat-tolerant varieties. In summary, AWD leads to lower panicle temperature and higher photosynthetic rate, sucrose content, and CTK level in panicles, which together enhance the heat tolerance at the PI stage, and therefore is a sustainable and feasible strategy to mitigate heat-induced yield loss in rice.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. Read at the source ↗

The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 17

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HDT significantly reduced rice yield in all four varieties except Shanyou 63.

Taxon
Oryza sativaOryza sativae1
Organ / stage
PI stagee2
Treatment
high daytime temperaturee2
Outcome
yielde0
Direction
reducede0
Qualifier
except for Shanyou 63e0
Comparator
control temperaturee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“HDT significantly reduced the yield of all four varieties except for Shanyou 63, primarily by decreasing the spikelet fertility and spikelet number”

✓ located verbatim in the source · abstract, characters 720–867 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK)”

✓ located verbatim in the source · abstract, characters 602–705 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HDT decreased spikelet fertility and spikelet number in rice.

Taxon
Oryza sativaOryza sativae1
Treatment
high daytime temperaturee2
Outcome
spikelet fertility and spikelet numbere0
Direction
decreasinge0
Comparator
control temperaturee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“HDT significantly reduced the yield of all four varieties except for Shanyou 63, primarily by decreasing the spikelet fertility and spikelet number”

✓ located verbatim in the source · abstract, characters 720–867 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK)”

✓ located verbatim in the source · abstract, characters 602–705 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD significantly improved spikelet fertility and spikelet number under HDT compared with continuous flooding.

Taxon
Oryza sativaOryza sativae1
Treatment
AWDe0
Outcome
spikelet fertility and spikelet numbere3
Direction
improvede0
Qualifier
under HDTe0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“while AWD significantly improved the two traits under HDT”

✓ located verbatim in the source · abstract, characters 869–926 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
e3

“HDT significantly reduced the yield of all four varieties except for Shanyou 63, primarily by decreasing the spikelet fertility and spikelet number”

✓ located verbatim in the source · abstract, characters 720–867 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HDT reduced the photosynthetic rate of rice.

Taxon
Oryza sativaOryza sativae1
Treatment
HDTe0
Outcome
photosynthetic ratee0
Direction
reducede0
Comparator
control temperaturee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“HDT reduced the photosynthetic rate, increased the starch content in leaves and stems, and decreased the transport of sucrose to panicles”

✓ located verbatim in the source · abstract, characters 938–1075 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK)”

✓ located verbatim in the source · abstract, characters 602–705 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HDT increased starch content in rice leaves and stems.

Taxon
Oryza sativaOryza sativae1
Organ / stage
leaves and stemse0
Treatment
HDTe0
Outcome
starch contente0
Direction
increasede0
Comparator
control temperaturee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“HDT reduced the photosynthetic rate, increased the starch content in leaves and stems, and decreased the transport of sucrose to panicles”

✓ located verbatim in the source · abstract, characters 938–1075 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK)”

✓ located verbatim in the source · abstract, characters 602–705 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HDT decreased sucrose transport to panicles in rice.

Taxon
Oryza sativaOryza sativae1
Organ / stage
paniclese0
Treatment
HDTe0
Outcome
transport of sucrosee0
Direction
decreasede0
Comparator
control temperaturee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“HDT reduced the photosynthetic rate, increased the starch content in leaves and stems, and decreased the transport of sucrose to panicles”

✓ located verbatim in the source · abstract, characters 938–1075 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK)”

✓ located verbatim in the source · abstract, characters 602–705 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD reduced panicle temperature in rice compared with continuous flooding.

Taxon
Oryza sativaOryza sativae1
Organ / stage
paniclese0
Treatment
AWDe0
Outcome
panicle temperaturee0
Direction
reducede0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“AWD reduced the panicle temperature, promoted sucrose synthesis in leaves, increased sucrose content and sucrose hydrolase activity in panicles under HDT”

✓ located verbatim in the source · abstract, characters 1077–1230 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD promoted sucrose synthesis in rice leaves compared with continuous flooding.

Taxon
Oryza sativaOryza sativae1
Organ / stage
leavese0
Treatment
AWDe0
Outcome
sucrose synthesis in leavese0
Direction
promotede0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“AWD reduced the panicle temperature, promoted sucrose synthesis in leaves, increased sucrose content and sucrose hydrolase activity in panicles under HDT”

✓ located verbatim in the source · abstract, characters 1077–1230 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD increased sucrose content and sucrose hydrolase activity in rice panicles under HDT.

Taxon
Oryza sativaOryza sativae1
Organ / stage
paniclese0
Treatment
AWDe0
Outcome
sucrose content and sucrose hydrolase activitye0
Direction
increasede0
Qualifier
under HDTe0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“AWD reduced the panicle temperature, promoted sucrose synthesis in leaves, increased sucrose content and sucrose hydrolase activity in panicles under HDT”

✓ located verbatim in the source · abstract, characters 1077–1230 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HDT increased abscisic acid content in rice panicles under continuous flooding.

Taxon
Oryza sativaOryza sativae1
Organ / stage
paniclese0
Treatment
HDTe0
Outcome
content of abscisic acide0
Direction
increasede0
Qualifier
under CFe0
Comparator
control temperaturee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“HDT also increased the content of abscisic acid and decreased that of cytokinins (CTKs), indole-3-acetic acid (IAA), and gibberellins (GAs) in panicles under CF”

✓ located verbatim in the source · abstract, characters 1232–1392 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK)”

✓ located verbatim in the source · abstract, characters 602–705 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HDT decreased cytokinins, IAA and gibberellins in rice panicles under continuous flooding.

Taxon
Oryza sativaOryza sativae1
Organ / stage
paniclese0
Treatment
HDTe0
Outcome
cytokinins (CTKs), indole-3-acetic acid (IAA), and gibberellins (GAs)e0
Direction
decreasede0
Qualifier
under CFe0
Comparator
control temperaturee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“HDT also increased the content of abscisic acid and decreased that of cytokinins (CTKs), indole-3-acetic acid (IAA), and gibberellins (GAs) in panicles under CF”

✓ located verbatim in the source · abstract, characters 1232–1392 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two temperature treatments at the PI stage (high daytime temperature, HDT; and control temperature, CK)”

✓ located verbatim in the source · abstract, characters 602–705 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD increased expression of CTK biosynthesis genes in roots and panicles under HDT.

Taxon
Oryza sativaOryza sativae1
Organ / stage
roots and paniclese0
Treatment
AWDe0
Gene
IPT , CYP735A , LOGe0
Outcome
expression of CTK biosynthesis genese0
Direction
increasede0
Qualifier
under HDTe0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“AWD increased the expression of CTK biosynthesis genes ( IPT , CYP735A , LOG ) and decreased that of CTK degradation gene CKX in roots and panicles under HDT”

✓ located verbatim in the source · abstract, characters 1394–1551 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD decreased expression of the CTK degradation gene CKX in roots and panicles under HDT.

Taxon
Oryza sativaOryza sativae1
Organ / stage
roots and paniclese0
Treatment
AWDe0
Gene
CKXe0
Outcome
expressione0
Direction
decreasede0
Qualifier
CTK degradation genee0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“AWD increased the expression of CTK biosynthesis genes ( IPT , CYP735A , LOG ) and decreased that of CTK degradation gene CKX in roots and panicles under HDT”

✓ located verbatim in the source · abstract, characters 1394–1551 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD enhanced xylem sap flow rate in rice.

Taxon
Oryza sativaOryza sativae1
Treatment
AWDe3
Outcome
xylem sap flow ratee0
Direction
enhancede0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“while enhanced the xylem sap flow rate and contents of CTKs, IAA, and GAs in panicles”

✓ located verbatim in the source · abstract, characters 1553–1638 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
e3

“AWD increased the expression of CTK biosynthesis genes ( IPT , CYP735A , LOG ) and decreased that of CTK degradation gene CKX in roots and panicles under HDT”

✓ located verbatim in the source · abstract, characters 1394–1551 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD enhanced contents of CTKs, IAA and GAs in rice panicles.

Taxon
Oryza sativaOryza sativae1
Organ / stage
paniclese0
Treatment
AWDe3
Outcome
contents of CTKs, IAA, and GAse0
Direction
enhancede0
Comparator
continuous flooding (CF)e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“while enhanced the xylem sap flow rate and contents of CTKs, IAA, and GAs in panicles”

✓ located verbatim in the source · abstract, characters 1553–1638 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
e2

“two irrigation treatments at the tillering stage including AWD and continuous flooding (CF)”

✓ located verbatim in the source · abstract, characters 506–597 · show in text ↑
e3

“AWD increased the expression of CTK biosynthesis genes ( IPT , CYP735A , LOG ) and decreased that of CTK degradation gene CKX in roots and panicles under HDT”

✓ located verbatim in the source · abstract, characters 1394–1551 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

AWD alleviated HDT damage more strongly in heat-sensitive than in heat-tolerant rice varieties.

Taxon
Oryza sativaOryza sativae1
Material
heat-sensitive varietiese0
Treatment
AWDe0
Outcome
alleviating effect on HDT damagee0
Direction
more pronouncede0
Comparator
heat-tolerant varietiese0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“AWD exhibited a more pronounced alleviating effect on HDT damage in heat-sensitive varieties than in heat-tolerant varieties”

✓ located verbatim in the source · abstract, characters 1653–1777 · show in text ↑
e1

“rice ( Oryza sativa L.)”

✓ located verbatim in the source · abstract, characters 356–379 · show in text ↑
Recommendationauthor interpretationoriginal resultpositivebasis n/a

Claim · model-written summary

AWD is a sustainable and feasible strategy to mitigate heat-induced yield loss in rice.

Outcome
heat-induced yield losse0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“AWD leads to lower panicle temperature and higher photosynthetic rate, sucrose content, and CTK level in panicles, which together enhance the heat tolerance at the PI stage, and therefore is a sustainable and feasible strategy to mitigate heat-induced yield loss in rice”

✓ located verbatim in the source · abstract, characters 1791–2061 · show in text ↑

irrigation · physiology: hormones · Oryza sativa (rice)Positive and negative results

4 papers: 3 positive, 1 negative or no effect

2 papers in the same direction as this paper · 1 reporting no difference from an alternative

  1. no effect
  2. positive
  3. positive
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processing · physiology: hormones · Oryza sativa (rice)Positive and negative results

8 papers: 7 positive, 1 negative or no effect

6 papers in the same direction as this paper · 1 in the opposite direction

  1. positive
  2. negative
  3. positive

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processing · molecular: expression · Oryza sativa (rice)Positive and negative results

11 papers: 9 positive, 2 negative or no effect

8 papers in the same direction as this paper · 2 in the opposite direction

  1. negative
  2. negative
  3. positive

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irrigation · molecular: metabolites · Oryza sativa (rice)Positive and negative results

55 papers: 44 positive, 11 negative or no effect

43 papers in the same direction as this paper · 9 in the opposite direction · 2 reporting no difference from an alternative

  1. positive
  2. negative
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processing · molecular: metabolites · Oryza sativa (rice)Positive and negative results

57 papers: 48 positive, 9 negative or no effect

47 papers in the same direction as this paper · 8 in the opposite direction · 1 reporting no difference from an alternative

  1. positive
  2. positive
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processing · yield · Oryza sativa (rice)Positive and negative results

114 papers: 88 positive, 26 negative or no effect

87 papers in the same direction as this paper · 21 in the opposite direction · 5 reporting no difference from an alternative

  1. positive
  2. positive
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irrigation · yield · Oryza sativa (rice)Positive and negative results

1,171 papers: 906 positive, 265 negative or no effect

905 papers in the same direction as this paper · 201 in the opposite direction · 64 reporting no difference from an alternative

  1. positive
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irrigation · molecular: expression · Oryza sativa (rice)Results agree

5 papers: 5 positive, 0 negative or no effect

4 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

1 paper more in this group

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8 of 10 groups shown: named compounds and organisms first, then groups whose results differ (checked claim by claim), then those with positive and negative results, then the smaller groups.

Claims are grouped by organism, treatment and what was measured or targeted. A paper counts as positive when any of its claims in the group is positive.