Paper · 2026 · Horticulturae

Biofilm of B. subtilis as a Growth Promoter of Lettuce (Lactuca sativa L.) in the Presence of Heavy Metals

Mirta Esther Galelli; Antonio Paz González; Ana Rosa García; et al.

14
claims
37
verbatim quotes
0
citations
3
research areas
✓
Open access · cc-by

Abstract

The negative effects of soil heavy metal contamination on food production could be mitigated using nature-based solutions, i.e., plant-growth-promoting rhizobacteria (PGPR). Yield of Lactuca sativa L. has been shown to increase by seed inoculation with biofilm of Bacillus subtilis subsp. spizizenii. The aim of this work was to assess whether this promoting effect occurs even in the presence of toxic concentrations of copper (Cu) and zinc (Zn). First, germination rates of lettuce seeds with increasing Cu and Zn concentrations were assessed. Then, lettuce plants growing from inoculated and non-inoculated seeds were cropped on substrates with excess Cu and Zn. Above- and below-ground lettuce variables were measured, and leaf macro- and microelements were determined. Germination was more severely affected by Cu overload than by Zn overload, while this trend was reversal for plant growth. Seed inoculation enhanced germination and increased plant growth assessed by root and shoot biomass, plant height and leaf area. For example, seed inoculation increased lettuce root and aerial biomass of lettuce growing on a metal- free substrate by 68% and 62%, respectively. This practice also promoted lettuce growth in metal-overloaded substrates, increasing aerial and root biomass by 32% and 29%, respectively, in connection with Cu, and by 260% and 183% when it came to Zn. Both Cu or Zn accumulated in the edible parts of lettuce growing on contaminated substrates, but seed inoculation did not mitigate metal uptake in any case. Except for Cu and Zn, macronutrients, micronutrients and heavy metal levels in lettuce leaves were affected neither by excess metal nor by seed inoculation. Altogether, B. subtilis biofilm has been proven to be an effective seed inoculant promoting seed germination and plant growth even in the presence of heavy metals. Adverse health effects due to metal accumulation in the lettuce edible parts are not expected to increase following seed inoculation.

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 14

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Cu overload affected lettuce seed germination more severely than Zn overload.

Taxon
lettuceLactuca sativae1
Treatment
Cu overloade0
Outcome
Germinatione0
Direction
more severely affectede0
Comparator
Zn overloade0

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“Germination was more severely affected by Cu overload than by Zn overload”

✓ located verbatim in the source · abstract, characters 774–847 · show in text ↑
e1

“lettuce seeds”

✓ located verbatim in the source · abstract, characters 476–489 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Zn overload affected lettuce plant growth more severely than Cu overload.

Taxon
lettuceLactuca sativae2
Treatment
Zn overloade1
Outcome
plant growthe0
Direction
reversale0
Comparator
Cu overloade1

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“this trend was reversal for plant growth”

✓ located verbatim in the source · abstract, characters 855–895 · show in text ↑
e1

“Germination was more severely affected by Cu overload than by Zn overload”

✓ located verbatim in the source · abstract, characters 774–847 · show in text ↑
e2

“lettuce plants”

✓ located verbatim in the source · abstract, characters 552–566 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Seed inoculation enhanced lettuce germination.

Taxon
lettuceLactuca sativae1
Treatment
Seed inoculatione0
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie2symbiont
Outcome
germinatione0
Direction
enhancede0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“Seed inoculation enhanced germination”

✓ located verbatim in the source · abstract, characters 897–934 · show in text ↑
e1

“lettuce seeds”

✓ located verbatim in the source · abstract, characters 476–489 · show in text ↑
e2

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Seed inoculation increased lettuce plant height and leaf area.

Taxon
lettuceLactuca sativae1
Treatment
Seed inoculatione0
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie2symbiont
Outcome
plant height and leaf areae0
Direction
increasede0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“Seed inoculation enhanced germination and increased plant growth assessed by root and shoot biomass, plant height and leaf area.”

✓ located verbatim in the source · abstract, characters 897–1025 · show in text ↑
e1

“lettuce plants”

✓ located verbatim in the source · abstract, characters 552–566 · show in text ↑
e2

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Inoculation increased lettuce root biomass by 68% without metal overload.

Taxon
lettuceLactuca sativae0
Treatment
seed inoculatione0
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie1symbiont
Outcome
root biomasse2
Direction
increasede0
Value
68%e0
Qualifier
metal- free substratee0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“seed inoculation increased lettuce root and aerial biomass of lettuce growing on a metal- free substrate by 68% and 62%, respectively.”

✓ located verbatim in the source · abstract, characters 1039–1173 · show in text ↑
e1

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
e2

“aerial and root biomass by 32%”

✓ located verbatim in the source · abstract, characters 1260–1290 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Inoculation increased lettuce aerial biomass by 62% without metal overload.

Taxon
lettuceLactuca sativae0
Treatment
seed inoculatione0
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie1symbiont
Outcome
aerial biomasse0
Direction
increasede0
Value
62%e0
Qualifier
metal- free substratee0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“seed inoculation increased lettuce root and aerial biomass of lettuce growing on a metal- free substrate by 68% and 62%, respectively.”

✓ located verbatim in the source · abstract, characters 1039–1173 · show in text ↑
e1

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Inoculation increased lettuce aerial biomass by 32% under excess Cu.

Taxon
lettuceLactuca sativae1
Treatment
This practicee1
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie2symbiont
Outcome
aerial biomasse3
Direction
increasinge0
Value
32%e0
Qualifier
Cue0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“increasing aerial and root biomass by 32% and 29%, respectively, in connection with Cu”

✓ located verbatim in the source · abstract, characters 1249–1335 · show in text ↑
e1

“This practice also promoted lettuce growth in metal-overloaded substrates”

✓ located verbatim in the source · abstract, characters 1174–1247 · show in text ↑
e2

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
e3

“root and aerial biomass of lettuce growing on a metal- free substrate”

✓ located verbatim in the source · abstract, characters 1074–1143 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Inoculation increased lettuce root biomass by 29% under excess Cu.

Taxon
lettuceLactuca sativae1
Treatment
This practicee1
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie2symbiont
Outcome
root biomasse0
Direction
increasinge0
Value
29%e0
Qualifier
Cue0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“increasing aerial and root biomass by 32% and 29%, respectively, in connection with Cu”

✓ located verbatim in the source · abstract, characters 1249–1335 · show in text ↑
e1

“This practice also promoted lettuce growth in metal-overloaded substrates”

✓ located verbatim in the source · abstract, characters 1174–1247 · show in text ↑
e2

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Inoculation increased lettuce aerial biomass by 260% under excess Zn.

Taxon
lettuceLactuca sativae1
Treatment
This practicee1
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie2symbiont
Outcome
aerial biomasse3
Direction
increasinge0
Value
260%e0
Qualifier
Zne0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“increasing aerial and root biomass by 32% and 29%, respectively, in connection with Cu, and by 260% and 183% when it came to Zn.”

✓ located verbatim in the source · abstract, characters 1249–1377 · show in text ↑
e1

“This practice also promoted lettuce growth in metal-overloaded substrates”

✓ located verbatim in the source · abstract, characters 1174–1247 · show in text ↑
e2

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
e3

“root and aerial biomass of lettuce growing on a metal- free substrate”

✓ located verbatim in the source · abstract, characters 1074–1143 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Inoculation increased lettuce root biomass by 183% under excess Zn.

Taxon
lettuceLactuca sativae1
Treatment
This practicee1
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie2symbiont
Outcome
root biomasse0
Direction
increasinge0
Value
183%e0
Qualifier
Zne0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“increasing aerial and root biomass by 32% and 29%, respectively, in connection with Cu, and by 260% and 183% when it came to Zn.”

✓ located verbatim in the source · abstract, characters 1249–1377 · show in text ↑
e1

“This practice also promoted lettuce growth in metal-overloaded substrates”

✓ located verbatim in the source · abstract, characters 1174–1247 · show in text ↑
e2

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Cu and Zn accumulated in edible lettuce parts on contaminated substrates.

Taxon
lettuceLactuca sativae0
Outcome
Cu or Zn accumulated in the edible partse0
Qualifier
contaminated substratese0

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“Both Cu or Zn accumulated in the edible parts of lettuce growing on contaminated substrates”

✓ located verbatim in the source · abstract, characters 1378–1469 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Seed inoculation did not mitigate lettuce metal uptake.

Taxon
lettuceLactuca sativae1
Treatment
seed inoculatione0
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie2symbiont
Outcome
metal uptakee0
Direction
did not mitigatee0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“seed inoculation did not mitigate metal uptake in any case”

✓ located verbatim in the source · abstract, characters 1475–1533 · show in text ↑
e1

“lettuce growing on contaminated substrates”

✓ located verbatim in the source · abstract, characters 1427–1469 · show in text ↑
e2

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Other macro- and micronutrient and heavy-metal levels in lettuce leaves were unaffected by excess metal.

Taxon
lettuceLactuca sativae0
Treatment
excess metale0
Outcome
macronutrients, micronutrients and heavy metal levels in lettuce leavese0
Direction
affected neithere0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“Except for Cu and Zn, macronutrients, micronutrients and heavy metal levels in lettuce leaves were affected neither by excess metal nor by seed inoculation.”

✓ located verbatim in the source · abstract, characters 1535–1691 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Other macro- and micronutrient and heavy-metal levels in lettuce leaves were unaffected by inoculation.

Taxon
lettuceLactuca sativae0
Treatment
seed inoculatione0
Interactor
Bacillus subtilis subsp. spizizeniiBacillus spizizeniie1symbiont
Outcome
macronutrients, micronutrients and heavy metal levels in lettuce leavese0
Direction
affected neithere0

Not reportedComparator

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

Lactuca sativa
Lactuca sativa · Taxonen:User:Geographer · CC BY 1.0 · Wikimedia Commons ↗

Verbatim

e0

“Except for Cu and Zn, macronutrients, micronutrients and heavy metal levels in lettuce leaves were affected neither by excess metal nor by seed inoculation.”

✓ located verbatim in the source · abstract, characters 1535–1691 · show in text ↑
e1

“biofilm of Bacillus subtilis subsp. spizizenii”

✓ located verbatim in the source · abstract, characters 253–299 · show in text ↑

copper atom · growth · Lactuca sativa (garden lettuce)Results differ

13 papers: 5 positive, 8 negative or no effect

7 papers in the same direction as this paper · 5 in the opposite direction

  1. positive
  2. positive
  3. positive

9 papers more in this group

See all 13 papers →

zinc atom · growth · Lactuca sativa (garden lettuce)Results differ

15 papers: 7 positive, 8 negative or no effect

7 papers in the same direction as this paper · 7 in the opposite direction

  1. positive
  2. positive
  3. positive

11 papers more in this group

See all 15 papers →

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.