Paper · 1989 · Evolution

SAFETY FACTORS IN VERTICAL STEMS: EVIDENCE FROMEQUISETUM HYEMALE

Karl J. Niklas

6
claims
13
verbatim quotes
11
citations
1
research areas
—
Not open access

Abstract

Predictions from a mechanical model for hollow vertical stems are tested against morphometric and mechanical studies of the vertical stems of Equisetum hyemale. The model predicts 1) that the wall thickness of hollow internodes must be at least 15% of the external radius of shoots, 2) that the elastic modulus of stems is quantitatively related to the ratio of apoplast (cell walls) to symplast (cytoplasm) areas in transverse sections through stems, and that (3) hollow stems are designed to sustain an additional and significant proportion of their own weight. The "safety factors" predicted for a hollow vertical stem are used to examine two adaptationist explanations for hollow stems: 1) "economy in design," which argues that natural selection will favor a reduction in the metabolic cost in constructing an organ, and 2) "mechanical design," which argues that stems are designed to maximize their mechanical stability during vertical growth. Evidence from E. hyemale indicates that 1) there is a developmental limit to the maximum allotment of biomass invested in the construction of stems, 2) as stem height increases, morphometric adjustments in internodal wall thickness occur which converge on predicted safety limits, and 3) the elastic modulus of stems changes as a function of the ratio of apoplast to symplast areas seen in transverse sections through shoots. Biomechanical and developmental evidence and the allometry of E. hyemale stems are consistent with the view that stems are designed for safety and are inconsistent with some predictions based on the economy in design.

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Claims 6

Observationhypothesisoriginal resultsimulated

Claim · model-written summary

The model predicts that the wall thickness of hollow internodes must be at least 15% of the external shoot radius.

Taxon
Equisetum hyemaleEquisetum hyemalee0
Outcome
wall thickness of hollow internodese1
Value
15% of the external radius of shootse1

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

Equisetum hyemale
Equisetum hyemale · TaxonDiego Delso · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“vertical stems of Equisetum hyemale”

✓ located verbatim in the source · abstract, characters 124–159 · show in text ↑
e1

“the wall thickness of hollow internodes must be at least 15% of the external radius of shoots”

✓ located verbatim in the source · abstract, characters 188–281 · show in text ↑
Observationhypothesisoriginal resultsignificant (as stated)simulated

Claim · model-written summary

The model predicts that hollow stems sustain an additional proportion of their own weight.

Taxon
Equisetum hyemaleEquisetum hyemalee0
Outcome
proportion of their own weighte2

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

Equisetum hyemale
Equisetum hyemale · TaxonDiego Delso · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“vertical stems of Equisetum hyemale”

✓ located verbatim in the source · abstract, characters 124–159 · show in text ↑
e2

“hollow stems are designed to sustain an additional and significant proportion of their own weight”

✓ located verbatim in the source · abstract, characters 465–562 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Internodal wall thickness is adjusted with increasing stem height, converging on predicted safety limits.

Taxon
Equisetum hyemaleEquisetum hyemalee0
Outcome
morphometric adjustments in internodal wall thicknesse3
Qualifier
as stem height increasese3

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

Equisetum hyemale
Equisetum hyemale · TaxonDiego Delso · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“vertical stems of Equisetum hyemale”

✓ located verbatim in the source · abstract, characters 124–159 · show in text ↑
e3

“as stem height increases, morphometric adjustments in internodal wall thickness occur which converge on predicted safety limits”

✓ located verbatim in the source · abstract, characters 1102–1229 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

The elastic modulus of stems changes with the ratio of apoplast to symplast areas in transverse sections.

Taxon
Equisetum hyemaleEquisetum hyemalee0
Organ / stage
shootse4
Outcome
elastic modulus of stemse4

Not reportedDirection

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

Equisetum hyemale
Equisetum hyemale · TaxonDiego Delso · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“vertical stems of Equisetum hyemale”

✓ located verbatim in the source · abstract, characters 124–159 · show in text ↑
e4

“the elastic modulus of stems changes as a function of the ratio of apoplast to symplast areas seen in transverse sections through shoots”

✓ located verbatim in the source · abstract, characters 1238–1374 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

There is a developmental limit to the maximum biomass allotted to stem construction.

Taxon
Equisetum hyemaleEquisetum hyemalee0
Outcome
maximum allotment of biomass invested in the construction of stemse5

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

Equisetum hyemale
Equisetum hyemale · TaxonDiego Delso · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“vertical stems of Equisetum hyemale”

✓ located verbatim in the source · abstract, characters 124–159 · show in text ↑
e5

“there is a developmental limit to the maximum allotment of biomass invested in the construction of stems”

✓ located verbatim in the source · abstract, characters 993–1097 · show in text ↑
Mechanismauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Hollow stems are designed for safety rather than economy in construction.

Taxon
Equisetum hyemaleEquisetum hyemalee0
Outcome
stems are designed for safetye6
Qualifier
Biomechanical and developmental evidencee7

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

Equisetum hyemale
Equisetum hyemale · TaxonDiego Delso · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“vertical stems of Equisetum hyemale”

✓ located verbatim in the source · abstract, characters 124–159 · show in text ↑
e6

“stems are designed for safety and are inconsistent with some predictions based on the economy in design”

✓ located verbatim in the source · abstract, characters 1489–1592 · show in text ↑
e7

“Biomechanical and developmental evidence and the allometry of E. hyemale stems”

✓ located verbatim in the source · abstract, characters 1376–1454 · show in text ↑