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Scaling the leaf length-times-width equation to predict total leaf area of shoots

Koyama Kohei Smith Duncan D. 帯広畜産大学

2022.06.08

概要

• Background and Aims An individual plant consists of different-sized shoots, each of which consists of
different-sized leaves. To predict plant-level physiological responses from the responses of individual leaves,
modelling this within-shoot leaf size variation is necessary. Within-plant leaf trait variation has been well investi-gated in canopy photosynthesis models but less so in plant allometry. Therefore, integration of these two different approaches is needed.
• Methods We focused on an established leaf-level relationship that the area of an individual leaf lamina is pro - portional to the product of its length and width. The geometric interpretation of this equation is that different-sized leaf laminas from a single species share the same basic form. Based on this shared basic form, we synthesized a new length-times-width equation predicting total shoot leaf area from the collective dimensions of leaves that comprise a shoot. Furthermore, we showed that several previously established empirical relationships, including the allometric relationships between total shoot leaf area, maximum individual leaf length within the shoot and total leaf number of the shoot, can be unified under the same geometric argument. We tested the model predic - tions using five species, all of which have simple leaves, selected from diverse taxa (Magnoliids, monocots and eudicots) and from different growth forms (trees, erect herbs and rosette herbs).
• Key Results For all five species, the length-times-width equation explained within-species variation of total
leaf area of a shoot with high accuracy ( R2 > 0.994). These strong relationships existed despite leaf dimensions scaling very differently between species. We also found good support for all derived predictions from the model (R2 > 0.85).
• Conclusions Our model can be incorporated to improve previous models of allometry that do not consider
within-shoot size variation of individual leaves, providing a cross-scale linkage between individual leaf-size vari-ation and shoot-size variation.

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APPENDIX

Derivation of eqn (8)

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15

16

Koyama & Smith — The length-times-width equation for shoot leaf area

shoot

shoot

By taking the product of both sides of the two lines in

eqn (14), and using eqn (2), we obtained:

mean (Aleaf ) ∝ max (Aleaf )

(15)

shoot

shoot

In deriving eqn (15), we assumed that the leaf that has maximum (or mean) length within each shoot also has the maximum (or mean) width in the same shoot. By multiplying both

sides of eqn (15) by the total number of leaves on each shoot

(N), we obtained:

N · mean (Aleaf ) ∝ N · max (Aleaf )

(16)

shoot

shoot

Because the left-hand-side of eqn (16) is Ashoot, we obtained

eqn (8).

Derivation of eqn (9)

In this section, we assume that lamina length and width of an

individual leaf are approximately proportional to each other

(Ogawa et al., 1995), and that individual leaf area can be predicted as the quadratic function of either leaf length or width

alone (Teobaldelli et al., 2019a, b):

Aleaf ∝ (Lleaf )

(17)

Equation (17) can also be used to predict the maximum Aleaf

within each shoot:

2

(18)

max (Aleaf ) ∝ max (Lleaf )

shoot

shoot

By multiplying both sides of eqn (18) by the total number of

leaves on each shoot (N), we obtained:

2

(19)

N · max (Aleaf ) ∝ N · max (Lleaf )

shoot

shoot

By combining eqns (8) and (19), we obtained:

2

(20)

Ashoot ∝ N · max (Lleaf )

By combining eqns (6) and (21), we obtained:

W f β+1 ∝ N · W f 2

(22)

Equation (22) can be solved for Wf:

W f ∝ N β−1

(23)

By substituting eqn (23) into eqn (6), we obtained:

β+1

(24)

Ashoot ∝ N β−1 ≡ N α

Derivation of eqn (10)

Equation (9) can be solved for Nf:

N ∝ Ashoot α

(25)

Using eqn (25), we obtained:

Ashoot

1− 1

∝ (Ashoot ) α ≡ (Ashoot )

(26)

Derivation of eqn (11)

By combining eqns (5) and (23), we obtained:

(27)

Lf ∝ W f β ∝ N β−1

By substituting eqn (27) into eqn (4), we obtained:

(28)

Ashoot ∝ W f · N β−1 ≡ W f · N γ

Derivation of eqn (13)

When different-sized foliage sets are affine to each other, the

maximum leaf size relative to the mean value and the minimum

leaf size relative to the mean value would be constants, independent of foliage size. This leads to the following relationship:

mean (Aleaf ) ∝ min (Aleaf )

(29)

shoot

shoot

By dividing both sides of eqns (15) and (29) by 2, and adding

them, we obtained:

shoot

By using our definition of Wf (eqn 3), eqn (20) can be rewritten as follows:

Ashoot ∝ N · W f 2

(21)

min (Aleaf ) + max (Aleaf )

shoot

(30)

mean (Aleaf ) ∝ shoot

shoot

By multiplying both sides of eqn (30) by N, we obtained

eqn (13).

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mean (Lleaf ) ∝ max (Lleaf )

shoot

shoot

(14)

mean (Wleaf ) ∝ max (Wleaf )

...

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