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Annals of Botany 90: 485±488, 2002

doi:10.1093/aob/mcf214, available online at www.aob.oupjournals.org

TECHNICAL NOTE

A Modern Tool for Classical Plant Growth Analysis


R . H U N T 1 , * , D . R . C A U S T O N 2 , B . S H IP L EY 3 and A . P . A S K E W 1
1Unitof Comparative Plant Ecology, Department of Animal and Plant Sciences, University of Shef®eld, Shef®eld S10
2TN, UK; 2Institute of Biological Sciences, Cledwyn Building, University of Wales, Aberystwyth SY23 3DD, UK and
3De partement de Biologie, Universite de Sherbrooke, Sherbrooke, Quebec, Canada J1K 2R1

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Received: 27 March 2002 Returned for revision: 21 May 2002 Accepted: 18 June 2002

We present an all-inclusive software tool for dealing with the essential core of mathematical and statistical cal-
culations in plant growth analysis. The tool calculates up to six of the most fundamental growth parameters
according to a purely `classical' approach across one harvest-interval. All of the estimates carry standard errors
and 95 % con®dence limits. The tool is written in Microsoftâ Excel 2000 and is available free of charge for use
in teaching and research from www.aob.oupjournals.org article supplementary data.
ã 2002 Annals of Botany Company

Key words: Relative growth rate, unit leaf rate, net assimilation rate, speci®c leaf area, leaf weight fraction, leaf area
ratio, allometry.

INTRODUCTION The tool derives relative growth rate in whole plant dry
Plant growth analysis is an explanatory, holistic and weight (RGR), the central parameter in plant growth
integrative approach to interpreting plant form and function. analysis, together with its components unit leaf rate
It uses simple primary data in the form of weights, areas, (ULR) (the `net assimilation rate', NAR, of many authors),
volumes and contents of plant components to investigate speci®c leaf area (SLA), and leaf weight fraction (LWF).
processes within and involving the whole plant (Evans, Instantaneously, these four are de®ned and related in the
1972; Causton and Venus, 1981; Hunt, 1990). From its following way:
origins at the end of the nineteenth century, plant growth
analysis ®rst illuminated plant physiology, then agronomy …1=W†…dW=dt† ˆ …1=LA †…dW=dt†  LA =LW  LW =W …1†
and now physiological and evolutionary plant ecology (e.g.
Garnier et al., 1999). In this journal alone, Hoffmann and RGR ULR SLA LWF
Poorter (2002) reported that 28 articles since 1993 have
drawn upon the approach in one way or another. where t is time, W is total dry weight per plant, LA is total
This paper presents an up-to-date and very comprehen- leaf area per plant and LW is total leaf dry weight per plant.
sive software tool that deals readily with the essential core Leaf weight fraction is synonymous with leaf weight ratio
of mathematical and statistical calculations in plant growth (LWR). The product of SLA and LWF, de®ned as LA/W and
analysis. These calculations can otherwise be tedious or, known as leaf area ratio (LAR), is also derived. These
particularly in the case of the statistics, neglected. growth parameters are generic and many analogues exist
We take the simplest possible approach, calculating the that accept other plant variables as inputs (see Hunt, 1990).
most fundamental of the growth parameters according to The purpose of our tool is to estimate all ®ve parameters,
purely `classical' methods across one harvest-interval including LAR, as mean values solely across one harvest-
(meaning the period of time between two successive interval (t1 to t2), with a standard error (s.e.) and 95 %
harvests). This contrasts with the `functional' or `dynamic' con®dence limits attached to each estimate. The root±shoot
approach, involving the use of many harvests and ®tted allometric coef®cient, and its s.e. and limits, are also derived
curves, which can be either parametric (Causton and Venus, for the same harvest-interval.
1981; Hunt, 1982) or form-free (Shipley and Hunt, 1996),
and also contrasts with the `combined' approach involving
curves ®tted to classically derived values (Poorter, 1989).
Here, we do not necessarily advocate the classical over the METHODS
functional approach; we simply facilitate the appropriate The mathematical and statistical methods follow those of
calculations for those who have already chosen the former Venus and Causton (1979), as developed by Causton and
for adequate reasons. Venus (1981), whereby harvest-interval means of the
various parameters are obtained without the use of ®tted
* For correspondence. Fax +44 (0)114 222 0002, e-mail r.hunt@shef. functions and without the `pairing' of plants across the
ac.uk harvest-interval.
ã 2002 Annals of Botany Company
486 Hunt et al. Ð A Modern Tool for Classical Plant Growth Analysis
The value of the exact mean RGR across the harvest- eqn (4) and as arrays of replicated measurements in
interval t1 to t2 is obtained from the usual formula by Fisher eqns (5) and (6). Using the contractions DW = W2 ± W1,
(1921): DLA = LA2 ± LA1, DlogeLA = logeLA2 ± logeLA1 and Dt
= t2 ± t1, the derivatives that appear in eqns (5) and (6)
R ˆ …loge W2 ÿ loge W1 †=…t2 ÿ t1 † …2† are:

¶E/¶W1 = ±DlogeLA/(DLA. Dt) (7)


and the statistical estimate of this quantity, RÃ, is the same.
From Causton and Venus (1981, eqn 2.33), the variance of R
is given by: @E=@W2 ˆ Dloge LA =…DLA :Dt†; …8†

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_…R† ˆ ‰_…loge W2 † ‡ _…loge W1 †Š=…t2 ÿ t1 †2 ; …3† @E=@LA1 ˆ f‰…ÿ1=LA1 †:DLA
‡Dloge LA Š:DWg=…DLA 2 :Dt†; …9†
where the symbol _ represents variance. In both of these
equations, W appears both at harvest 1 and at harvest 2: in
eqn (2) it appears as harvest-means of logeW (not as logs of @E=@LA2 ˆ f‰…1=LA2 †:DLA ÿ
harvest means of W; see Hoffmann and Poorter, 2002), and Dloge LA Š:DWg=…DLA 2 :Dt†; …10†
in eqn (3) it appears as arrays of replicated measurements.
The degrees of freedom associated with the estimate of
RGR, and with all the other parameters except the @ 2 E=@W1 2 ˆ @ 2 E=@W2 2 ˆ 0; …11†
allometric coef®cient, are n ± 2, where n is the total number
of plants for both harvests combined.
For mean ULR (NAR) across the same harvest-interval, @ 2 E=@LA1 2 ˆ f‰…1=LA1 2 †:DLA 2 ÿ …2=LA1 †:DLA
the approximate value is obtained from the usual formula by ‡2:Dloge LA Š:DWg=…DLA 3 :Dt†; …12†
Williams (1946):

E  ‰…W2 ÿ W1 †…loge LA2 ÿ loge LA1 †Š= @ 2 E=@LA2 2 ˆ f‰…ÿ1=LA2 2 †:DLA 2 ÿ …2=LA2 †:DLA ‡
‰…LA2 ÿ LA1 †…t2 ÿ t1 †Š; …4† 2:Dloge LA Š:DWg=…DLA 3 :Dt†; …13†

where the symbol » means `is approximately equal to'. The @ 2 E=…@W1 @LA1 † ˆ ‰…1=LA1 †:DLA ÿ
statistical estimate of unit leaf rate, EÃ, was derived by Dloge LA Š=…DLA 2 :Dt†; …14†
Causton and Venus (1981, eqn 2.34) and is:

Eà  E ‡ ÿ
1 2 2
2 …@ E=@W 2 †: _ …W2 †
@ 2 E=…@W2 @LA2 † ˆ ‰…1=LA2 †:DLA ÿ
1
‡ÿ 2 2
2 …@ E=@W1 †: _ …W1 †
Dloge LA Š=…DLA 2 :Dt†: …15†
1 2 2 1 2 2
‡ÿ
2 …@ E=@LA2 †: _ …LA2 † ‡ÿ
2 …@ E=@LA1 †: _ …LA1 †
2
For the LAR, the quotient at either harvest is de®ned
‡‰@ E=…@W2 @LA2 †Š:C…W2 ; LA2 †‡ simply as f = LA/W ((Briggs et al., 1920). The statistical
‡‰@ 2 E=…@W1 @LA1 †:C…W1 ; LA1 † …5† estimate of f, and its variance, both after Causton and Venus
(1981, eqns 2.13 and 2.14), are then given by:

where C represents a covariance term. The variance of this


expected mean ULR, again from Causton and Venus (1981, fà  exp‰loge f ‡ ÿ
1
2 _ …loge f †Š …16†
eqn 2.35), is given by:
and
2 2
_…E†  …@E=@W2 † : _ …W2 † ‡ …@E=@W1 † : _ …W1 †
_ (f) » exp[2 loge f + _(loge f)]. {exp[_(loge f)] ± 1} (17)
‡…@E=@LA1 †2 : _ …LA1 † ‡ …@E=@LA2 †2 : _ …LA2 †
‡2 …@E=@W2 †…@E=@LA 2 †:C…W2 ; LA2 † with replicated arrays of quotients being used in eqn (17).
‡2 …@E=@W1 †…@E=@LA1 †:C…W1 ; LA1 †: …6† Having just two harvests provides no information about the
exact relation between f and time, so, from the separate
estimates of f on each harvest occasion, the assumed mean
Again, variables W and LA appear both at harvest 1 and value of LAR across the harvest-interval t2 ± t1, and its
harvest 2: as harvest-means of W, LA and loge LA in variance, are simply
Hunt et al. Ð A Modern Tool for Classical Plant Growth Analysis 487
1 charge to those with access rights, from the Annals of
F ˆ …fÃ1 ‡ fÃ2 † …18†
2 Botany's own Internet archive at www.aob.oupjournals.org
article supplementary data.
and
Inputs
_(F) = 12 [_(f1) + _(f2)] (19)
The input matrix is arranged in the usual statistical
Means of SLA and LWF (Evans and Hughes, 1961), and format, i.e. columns for variables and rows for cases, which
their variances, are obtained by methods that parallel those are the individual plants. Maximally, the variables are time
used for LAR. of harvesting, total root weight per plant, total leaf weight
The root-shoot allometric coef®cient (the slope of per plant, total non-leaf above-ground weight per plant (all
weights to be expressed uniformly on a dry- or fresh-weight

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logeWroot on logeWshoot) and its variance are obtained
using a bivariate maximum likelihood estimation (Causton basis), and total leaf area per plant. Not all of these variables
and Venus, 1981, eqns 6.28, 6.29, 6.32 and 6.48; see also are essential (see below).
Sprent, 1969, Chapter 3). With x as logeWshoot and y as
logeWroot, and using the contractions Outputs
2 2
Sxx ˆ S…x ÿ x† ; Syy ˆ S…y ÿ y† ; and These are mean values for the speci®ed harvest-interval
Sxy ˆ S…x ÿ x†…y ÿ y†; of each of RGR, ULR (NAR), LAR, SLA, LWF and the
root±shoot allometric coef®cient, all with s.e. and a 95 %
the maximum likelihood estimate of the coef®cient of the con®dence limit attached. Figure 1 shows the tool display-
slope of the straight line joining the two bivariate sets of ing a specimen set of input and output data with n = 24.
data is given by:
 q
Units
à ˆ …Syy ÿ :Sxx † ‡ …Syy ÿ :Sxx †2 ‡ 4:S2xy =2:Sxy …20†
For each of the dimensions time, weight and area, and in
both the input and the output ®elds, the user selects units
where, assuming that the covariances of the bivariate from extensive drop-down menus. In any or all of these
normal distributions are zero, dimensions, the units selected for the outputs may differ
from those used for the inputs (the tool will perform the
 ˆ ‰_…y1 † = _ …x1 † ‡ _…y2 † = _ …x2 †Š=2 …21†
appropriate conversions). Most of the units offered are
metric, but some are non-SI. It is up to the user to make
The intercept term is estimated by: sensible choices according to the nature of the problem in
à hand and the eventual use to which the outputs will be put.
aà ˆ y ÿ x …22†

and, since for i = 1, 2


` F R E Q U E N TL Y A S K E D Q U E S T I O N S '
à i ÿ aà †=…
Ãi ˆ …xi ‡ y à ‡ Ã2 †; …23† Does it matter how far apart the harvests are spaced? No,
the time interval presented by the user will be the time
then the variance of the slope is given by: interval used for all calculations. Of course, the larger the
time interval between harvests, the more the average values
_…b† ˆ f‰_…x1 † ‡ _…x2 †Š=2g=S…Ãi ÿ †2 …24† of the growth parameters will differ from the instantaneous
values if the plants are not growing exponentially.
The degrees of freedom associated with the estimate of the What if I have more than two harvests? Only two harvests
allometric coef®cient are n ± 4. will be recognized from the input data: those occurring at
the proffered tmin and tmax. Any data presented with
intermediate values of t will be ignored, so it is best to
T H E SO F T W A R E T O O L present only two harvests at a time, re-using the tool for
each different harvest-interval, species and treatment com-
In outline
bination. More elaborate methods of analysis involving
The tool runs on a PC and is written in Microsoftâ Excel curve-®tting might be indicated if harvests form an appre-
2000. The tool activates every time a suitable matrix of data ciable series (see Causton and Venus, 1981; Hunt, 1982;
is copy-pasted into the input ®eld (in `values only' mode). Shipley and Hunt, 1996).
The outputs comprise a complete set of growth parameters Does it matter what units are chosen? No (see above).
for the one harvest-interval, all with standard errors and However, the units chosen must be consistent within any
95 % limits. The tool is liberally supplied with drop-down one variable throughout any one use of the tool.
comments explaining and advising on each part of the Do there have to be equal numbers of plants at each
procedure. The tool is available free of charge from R.H. or harvest? No, but sparse or unbalanced replication will
B.S. Alternatively, it may be downloaded, also free of adversely affect the statistical results.
488 Hunt et al. Ð A Modern Tool for Classical Plant Growth Analysis
modi®ed form. For example, without Wroot only SLA will be
delivered unaltered; RGR, ULR, LAR and LWF will all be
calculated with Wshoot standing-in alone for total weight W.
This facility creates the possibility of deliberate alternative
analyses. The only invariably essential datum is time.
What if I want to calculate growth parameters quite other
than the ®ve already included? Because the tool is generic,
by manipulating the input variables the user can calculate
any other growth parameters that have the same mathemat-
ical and statistical structure as RGR, ULR or LAR. In such
cases the inputs and outputs will not be labelled correctly,
but this can be allowed for.

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ACKNOWLEDGEMENTS
Peter Mitchell and Eric Garnier commented very helpfully
on a draft of this manuscript; Bill Hoffmann kindly allowed
us to cite work then in press.

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Evans GC. 1972. The quantitative analysis of plant growth. Oxford:
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F I G . 1. The spreadsheet tool for classical plant growth analysis,
Effect of arti®cial shading upon Impatiens parvi¯ora. New
displaying a specimen set of input and output data. The tool is available
Phytologist 60: 150±180.
free of charge from www.aob.oupjournals.org article supplementary data.
Fisher RA. 1921. Some remarks on the methods formulated in a recent
article on `The quantitative analysis of plant growth'. Annals of
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What are the lower and upper limits for numbers of structure: an ecophysiological perspective. New Phytologist (special
replicate plants? Two plants minimum per harvest; ®ve issue) 143: 1±221.
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relative growth rate. Annals of Botany 90: 37±42.
degrees of freedom for the allometric coef®cient are n ± 4); Hunt R. 1982. Plant growth curves: the functional approach to plant
100 plants maximum for both harvests combined. growth analysis. London: Edward Arnold.
What if a variable contains missing values? The tool will Hunt R. 1990. Basic growth analysis. London: Unwin Hyman.
disregard any cases (i.e. rows or single plants) having one or Poorter H. 1989. Plant growth analysis: towards a synthesis of the
more missing values (empty cells). Missing values will not classical and the functional approach. Physiologia Plantarum 75:
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be equated to zero. Cases can also be deleted temporarily to Shipley B, Hunt R. 1996. Regression smoothers for estimating parameters
investigate the effect of eliminating potential data outliers. of growth analyses. Annals of Botany 76: 569±576.
What if one whole variable is missing throughout? Fill Sprent P. 1969. Models in regression, and related topics. London:
this variable's range with zeros. The calculations will Methuen.
proceed wherever possible, but will omit any parameters Venus JC, Causton DR. 1979. Plant growth analysis: a re-examination of
the methods of calculation of relative growth and net assimilation
requiring the missing variable(s), e.g. without LA only RGR, rates without using ®tted functions. Annals of Botany 43: 633±638.
LWF and the allometric coef®cient will be calculated. In Williams RF. 1946. The physiology of plant growth with special reference
certain instances some calculations will proceed, but in to the concept of net assimilation rate. Annals of Botany 10: 41±72.

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