103

A QUANTITATIVE: ANALYSIS O F PLANT GR,OWTH.

PART I.

BY G. E. BRIGGS, M.A. (CANTAB.),

E’RANKLIN KIDD, M.A. ( CANTAB.), D.Sc. (LoND.),
AND

CYRIL WEST, A.R.C.Sc., D.Sc. (LoND.).

(With 9 text-figures.)
PAQE
INTRODUCTION . . . . . . 103 . . .
I. The Relative Growth-Rate Curve for Maize 106
CHAPTER
AverageGrowth-Rate . .
120 . .
Summary . . . .
120 . .
INTRODUCTION.
THE quantitative analysis of plant growth is a branch of plant physiology
to which adequate attention has not as yet been paid, but which should
be able nevertheless t o yield results of much theoretical interest and
economic importance. Methods for obtaining data for the analysis of
plant growth under ordinary cultural conditions are in general simple,
consisting principally of periodic dry-weight and leaf-area measurements,
and a quantity of excellent data of this nature has already been collected
and exists in the literature. A5 yet a thorough analysis of these results
has not been presented. Attempts have been made to fit in a few isolated
results with various empirical laws without wide examination of existing
data.
For example i t has been recently suggested by V. H. Blackman(1)
that the growth of an annual plant can be treated as a process following
the compound interest law expressed by the formula
W=
where W = the dry-weight of the plant a t time t , W,, = the initial dry
weight of the plant, r = the rate of interest or “efficiency index” of dry-
weight production, and e = the base of the natural logarithms.
104 Quantitative Analysis of P l a i ~ tGrowth
Another suggestion is that the growth of a plant is similar to an
autocatalytic reaction, and that it can be expressed by the formula

where A = the maximum dry-weight of the plant, x = the dry-weight

of the plant a t any time. t , t, = the time at which the weight of the
plant is half the final dry-weight, and K = a constant. This suggestion
was put forward by Robertson (24 and 25) and has received the support
of Reed and Holland(a1) and of Rippelcn and 23).
Finally Mitscherlich(14, 15 and 16) has attempted to apply to plant
growth as measured by dry-weight increase the following formula
= log ?A- c . 2.
log ( V A- 7;)
I n this formula n = a variable quantity indicating the probable
number of environmental factors, A = the maximum possible dry; weight
attainable by the plant in question, y = the dry-weight of the.plant at
time x, the time x being expressed in vegetation periods (Vegetations-
abschnitten) of arbitrary length.
A fuller consideration and criticism of these suggestions will be given
in subsequent chapters.
I n the present paper the primary objective is to attempt to obtain
a concrete idea of the growth and development of the plant. At the
outset i t will be best to confine our attention to simple cases and we
propose t o devote the first chapter to a consideration of an annual plant.
Certain data are required : periodic dry-weight measurements of the
whole plant (and its various parts) a t short intervals throughout its life,
starting from the seed a t the time of sowing; corresponding periodic
leaf-area measurements; data with regard to light, temperature, and
water supply. To avoid the error due to individual variation, a large
number of plants should be used for each dry-weight measurement and
where possible uniform ‘pure-line’ material should be employed.
There are various methods of presenting the results, and in the first
instance we shall use the relntive growth-rate curve. The principle of the
proposed method of expressing rate of growth is analogous to that of
the method by which the rate of most reactions, both chemical and
physiological, are expressed, namely, amount of change per unit of
material per unit of time. Since the amount of material in the growing
plant is constantly changing, and since the relative rate of growth is
not constant, as the following analysis will show, to achieve mathe-
*
 G. E. BRIGGS,
F. KIDD,AND C. WEST 105
matical accuracy the increase should be measured over an infinitely
short period. This procedure is manifestly impossible, and as we have
no exact knowledge of the way in which the relative rate of growth
varies over a given period we have adopted the following purely con-
ventional method of defining relative rate of growth, The relative rate
of growth of a plant during any given week in its life-cycle is the amount
Relative growth-rate curves for " Badischer Friih " Maize, 1876

Week from sowing

Fig. 1.

of dry matter which 100 g. of dry matter taken a t the beginning of the
week adds during the week. A week has been chosen since this i3 the
usual interval between determinations of dry-weight in most experi-
ments on growth in plantsl. It must be realised that the method does
not pretend t o mathematical accuracy being merely an approximate
average for the week, but with such results as are a t present available
nothing more accurate can be obtained. Even if measurements over

When results are not given for a week we have calculated the increase per 100 g.
for the period and divided the result by the number of weeks in the period: for example,
if the period is 8 days and 40 g. increases by 20 g. during t h a t period, then therelative
-
rate is
20 x 100 . 8
40
--
' 7 '
106 Quuntitative Amlysis of Plant Grawth
shorter intervals were available, until we gain knowledge of a mathe.
matical law according to which the rate changes, we cannot determine
the rate at any given time.
It might be suggested that allowance could easily be made for the
continuous increase in the dry-weight during the week by assuming that
this takes place a t a uniform rate, and consequently that by means of
the following logarithmic formula the rate could be determined:
log w - log w,= r,
where W = the dry-weight at the end of the week, and W , = the dry-
weight a t the beginning of the week.
I n curve A, Fig. 1, this allowance has been made. In Curve B the
ordinates are relative growth-rates calculated by our method, that is,
without making allowance for the continuous increase during the week.
These curves show similar variations in relative rate from week to week.
The more complicated method, however, does not achieve accuracy as
it rests on the assumption that the rate remains constant during the
week, an assumption manifestly incorrect since the rate varies from
week to week. Both methods me purely conventional and only approxi-
mate t o accuracy, and nothing definite is t o be gained by adopting the
more complicated procedure.
The'relative rate of plant growth a t any time may be taken as an
expression of the efficiency of the plant a t that time in producing dry
matter. It must be remembered from what we have said above that
the actual value of the figures for the growth-rate is only an average of
the changing rate during a week. They are, however, valid for purposes
of comparing the rate of a plant's growth from week to week.
The gist of the method described above of presenting the results of
growth experiments has been previously briefly put forward by Kidd
and West (9).

CHAPTER I.

THE RELATIVE GROWTH-RATE CURVE FOR MAIZE.

The most complete set of data for one plant is to be found in a series
of papers published in Germany many years ago under the general
<directionof U. Kreusler(l0, 11, and 13). From among the many results
recorded, we have chosen those for maize, since the growth of this plant
WRS studied in four successive years. The data include not only weekly
dry-weight measurements and corresponding leaf-area measurements,
 C. F,. RRIGGS,F. KIDD, AND C. WEST 107
but also environmental conditions such as light, temperature, water-
supply, etc. The dates of the first appearance of the flowers and of seed
formation are also given. The work appears to have been carried out
without any pre-conceived idea as to what the results would be, and the
results themselves have not as yet been worked out nor have they re-
ceived critical consideration although collected and published 40 years
ago. These results will be analysed in this and in the following chapter,
and certain interesting conclusions reached. We have constructed from
Kreusler’s data the tables and figures presented in this paper. Figs. 2
and 3 show respectively the relative growth-rate curves for “ Badischer
Friih” maize, the rates being calculated, as above described, on the
basis of weekly periods for the years 1875-1878 inclusive, and for five
different varieties of maize calculated on the same basis for the year 1875.

Table I.-“ Badischer Fruh )) Maize grown at Poppelsdorf in 1875.

Total dry Increase in Weekly Ratio of Record of
weight dry-weight percentaee leaf-area Mean a pearance
Date of Growth of a single since last increase i n Leaf- tempera- o f d and 0
harvest period plant harvest dry-weight area $e$& ture flowers
since last Sq. cm. Sq. cm.
1875 Days Gm. Om. harvest per plant per gm. “C.

11th May 0.206

( 3 ~ *
1st June 21 0.268 0.062 10 33.3 124 14
8th ,, 7 0.559 0.291 108 97.8 177 19.3
15th ,, 7 1.069 0.510 91 181.1 170 17-1
(129)
23rd ,, 8 2.448 1.379 113 405.1 167 166
30th ,, 7 4.776 2.328 95 889 186 17.0
7th July 7 11.077 6.301 132 1543 139 19.9 dflowers
(113)
13th ,, 6 23419 12.542 132 2646 112 18.1 ?flowera
(85)
21~t $, 8 43.844 20.225 74 3633 83 18.8
(28)
27th ,, 6 55.934 12.090 33 3291 59 17.6
3rd Aug. 7 72.875 16.941 30 3617 50 16.9
10th ,, 7 76.619 3.744 5 3630 48.5 19.1
17th ,, 7 84.332 7.713 10 2933 34.5 215
24th ,, 7 89421 5.289 6 2696 30 19.8
31st ,, 7 100.380 10.759 12 2907 29 19.0
7th Sept. 7 130.478 30.098 30 2986 23 16.1
(21)
15th ,, 8 158.139 27.661 18 2335 14.8 17.5

* The figures in brackets in column 5 give the percentage increase in dry-weight for
he number of days stated in column 2.
 108 Quantitative Analysis of Plapat Growth
Table 11.--" Badischer Fruh " Maize grown at Poppelsdorf in 1876.
Total dry- Increase in Weekly Katio of Number M&n Record$
wei ht of dryweight ercentage leaf area of hours tempera- appeuraia
Date of Growth a skngle since last kcrease in Leaf- to dry- of sun- ture for of $ and 0
harvest period plant harvest dry weight area weight shine the week Rowen
sincelast Sq cm Sq cm.
18'76 Days Gm. Gm harvest per plant per gm C
11th May 0.3264
( - 2.91)*
24th ,, 13 0.3169 -0.0095 -1.57 9.8
31st ,, 7 0.2724 -0,0445 -14 8.4 31 42 13
7th June 7 0.2914 fO.0190 +7 19.0 65 61 15.1
14th ,, 7 0.3642 0.0728 25 41.4 113 16 16.3
21st ,, 7 0.5674 0.2032 56 92.0 162 57 16.9
28th ,, 7 2.0733 1-5059 260 350.8 170 102 19
5th July 7 5.655 3.582 164 987.0 172 42 17.6
12th ,, 7 11,151 5.496 97 1794.5 159 45 20
19th ,, 7 30.265 19.114 170 3272.6 108 71 17.6 J s n d v
26th ,, 7 58.609 28.344 93 49594 85 49 18.5 flowers
2nd Aug. 7 106.908 48.299 83 6196.7 58 93 20.3
9th ,, 7 131.169 24.261 22.7 6530.7 42 76 18-3
16th ,, 7 207.373 76.204 58.2 6666.7 32 94 21.9
23rd ,, 7 204.436 -2.937 - 1.42 6201'4 30.5 77 21.6
30th ,, 7 202'168 -2.268 -1.12 4231'4 21 10 14.5
* The figure in brackets in column 5 gives the percentage increase in dry-weight for 13 daya

Table 111.-"Badischer Fruh" Maize grown at Poppelsdorf in 1877.

Total dry- Increase in Weekly Ratio of Number Mean Recordd
weight of dry-weight percentage leaf-area of hours tempem- appeanna
Date of Growth a single since hat increase in Leaf- to dry- of sun- ture for of .jand 9
harvest period plant harvest dry weight area weight shine the week Bowem
since last Sq. cm. Sq. cm.
1811 Days Om. om. harvest perplant pergm. 'C.
17th May 0.3353
( - 3.88)*
29th ,, 12 0.3223 -0.013 - 1.9 14 11.9
5th June 7 0.2819 -0404 -12.6 5.43 19.2 44 16.6
12th ,, 7 0.2877 +OW58 f 2.13 45.7 159 33 19.1
19th ,, 7 0.9395 0.6518 227 168.8 180 54 18.7
26th ,, 7 2.500 1.650 176 477.5 192 32 18.8
3rd July 7 6.365 3.775 150 1060 166 40 18.0
10th ,, 7 10.637 4.272 67 1671 157 20 14.7
17th ,, 7 24.447 13.810 130 3216 132 27 19.0 dflowen
24th ,, 7 41.408 16-961 69 3788 92 38 17.5 ?flowers
31st ,, 7 66.498 25.09 61 4591 69 20 18.6
7th Aug. 7 88.654 22.156 33.4 4934 55.5 40 16.4
14th ,, 7 119.842 31.188 35 5298 44 19 19.3
21st ,, 7 135.532 15.690 13 4852 35.5 30 19.5
28th ,, 7 140.782 6.250 3.9 4158 29-5 17 18.1
4thSept. 7 179.973 39.191 28 4332 23 25 16.6
11th ,, 7 187.795 7.822 4.35 4035 21.5 17 13.0
18th ,, 7 201.293 13.498 7.2 18 16.0
25th ,, 7 220.709 19.416 9.4 9 9.7
2nd Oct. 7 199.970 -20.739 - 9.4 22 7.9
9th 1, 7 204.017 +4*047 + 2.0 9 9.0
* The figure in brackets in column 5 gives the percentage increase in dry-weight for 12 days
 G. E. BRIWS,F. KIDD,AND C. WEST 109
Table IV.--" Badischer Friih " Maize g r m n at Poppelsdorf in 1878.
Total dry- Increase in Weekly Ratio of Number Record Of
wei ht of dry-weight led-uea of hours Yean a peaonce
Date of
hV&
Growth
period
a skgle
plsnt
since lsst
harveat
E23:
dry-weight
Leaf-
area
to dr
weigKi
of8UD- temperr-
shine ture
o r b and 0
flowers
since lsst Sp. cm. Sq. cm.
1818 Days Gm. Gm. harvest per plsut per gm. 'C.
20th May 0.3282
28th ,, 8 0.3280 - 0.0002 12.4
4th June 7 0-2870 - 0.041 - 12-5 40 13-6
11th ,, 7 0.2550 - 0'032 -11.2 17.9 70 27 15-5
18th ,, 7 0.3080 + 0.053 f20.8 29.2 95 19 15.1
25th ,, 7 0.6370 0.329 106.5 124.4 195 40 17.9
2nd July 7 2.319 1.682 264 419.2 181 36 19.8
9th ,, 7 4.654 2.335 100 762-2 174 16 17.1
16th ,, 7 9.019 4.365 94 1301 144 20 16-8
23rd ,, 7 20401 10.982 122 2136 107 57 19.6 ZandQ
30th ,, 7 34.557 14.556 72 2805 81 23 19.8 flowers
6th Aug. 7 57.587 23.030 66 3384 59 35 18.2
13th ,, 7 70.095 12.058 21.7 3047 43.5 32 20.1
20th ,, 7 85.165 15.070 21.4 3025 35.5 35 19.3
27th ,, 7 111.649 26.484 31 2976 26.5 17 17.8
3rd Sept. 7 124.760 13.111 11.7 2684 21.5 21 19.0
10th ,, 7 121.990 - 2.770 -2.2 2387 19.5 35 19.2

Table V.--" Huhner " Maize grown at Poppelsdorf in 1875.

Total d r y Increase in Weekly asti0 of Becord of
weight of dry-weight leaf-area Mean appesnnce
Date of Growth a single since last ~~~~~~~ Leaf- to dry- tempers- of d and 0
harvest period plant harveat dry-weight are8 weight ture Bowers
sincelsat Sq.cm. &.om.
1815 Days Om. Om. harvest perplant pergm. 'C.
11th May 0.127
(17)*
1st June 21 0.149 -022 5.7 28.9 194 14.0
8th 9 s 7 0.476 ,327 220 86-3 181 19.3
15th ,, 7 0.824 -348 73 153 186 17.1
( 114)
23rd ,, 8 1.765 -941 100 371 210 16.6
30th ,, 7 24347 1.082 61 627 220 17.0
7th July 7 7.292 4.445 156 895 123 19.9 8 and$?
(59) flowers
13th ,, 6 11.570 4.278 69 749 65 18.1
(86)
21st. ,, 8 21676 10-006 75 1410 66 18-8
(89)
27th ,, 6 40.735 19.159 104 2126 52 17.6
3rd Aug. 7 55.918 15.183 37 1917 34 16.9
10th ,, 7 60.648 4.730 8.5 2196 36 19.1
17th ,, 7 73-946 13.298 22 2254 31 21.5
24th ,, 7 90.491 16.545 22 2090 23 19.8
31st ,, 7 88.212 -2.279 -2.5 2032 23 19.0
7th Sept. 7 81.618 -6.594 -7.5 1367 17 16.1
( - 6.4)
15th ,, 8 76.385 -5:233 -5.6 665 9 17.5
*' The figures in bracketa in column 5 give the percentage increase in dry-weight for
the number of days stated in column 2.
110 Quantitative Analysis of Plant e o w t h
Table VI.--" 0berl;iinder" Maize gtbwn at Poppelsdorf in 1815.
Total dry. Increase in Weekly Ratio of Record of '
weight of dry-weight percentage leaf-sres Mean appearance
Date of Growth a single since last increase in Lest tempera- of 6 and 0
harvest period plant harvest dry-weight area &!:it ture flowers
since last Sq. cm. Sq.em.
1875 Days Gm. Gm. harvmt per plant pergm., "C.
11th May 0.107
(51I*
1st June 21 0.162 0.055 17 31 192 14.0
8th ,, 7 0.467 0.304 188 96.7 207 19.3
15th ,, 7 0.955 0488 105 165.3 173 17.1
23rd ,, 8 1.839 0.884 374.1 203 16.6
30th 7 3,097 1.258 67 621.0 201 17.0
7th JZy 7 6-758 3.661 118 668.6 99 19.9 J and 0
flowers
13th ,, 6 14.448 7.690 ):::( 9506 66 18.1
21st 8 24.890 10.442 (2 1098 44 18.8
27th 6 30.180 5.290 (2215) 1407 47 17.6
3rd A&. 7 51.196 21.016 70 1653 32 16.9
10th ,, 7 70.978 19.782 39 2217 31 19.1
17th ,, 7 59.110 -11.868 -17 2359 40 21.5
24th ,, 7 81.687 22.577 38 1464 18 19.8
31st ,, 7 73.921 -7.766 -9 1235 16 19.0
7thSept. 7 74,120 0.199 0.3 1716 10 16.1
15th ,, 8 87.192 13.072 514 (:58)6 17.5
* The figures in brackets in column 5 give the percentage increase in dry-weight for
the number of days stated in column 2.

Table VI1.-" Ungarischer Friih " Maize grown at Poppelsdorf in 1875.

Total dry- Increase in Weekly Ratio of Fkcord of
weight of dry-weight percentage leaf-area Mean appearance
Date of Growth n single uince last increase in Leaf- to dry- tempera- of J and 0
harveat period plant harvest dry-weight area weight ture flowers
since last Sq.cm 5q.cm.
1875 Days Gm Gm harveat per plant pergm. "C.
11th May 0.2696
( - 0.88)'
1st June 21 0.2672 - 04024 -0.29 31.0 116 14.0
8th ,, 7 0.609 0.342 128 118.9 196 19.3
15th ,, 7 1.110 0.501 82 190.2 172 17.1
(93)
23rd
30th
., 8
7
2.143
4.123 '
1.033
1.980
81
93
393.2
807.0
184
196
16.6
17.0
7th J& 7 12.101 7.978 195 2109 174 19.9 6 and?
(92) flowers
13th .. 6 23.244 11.143 107 3030 130 18.1
(91)
21st ,, 8 44.48 21.236 80 4329 97 18.8
27th 6 70.46 25.98 65 5635 80 17.6
3rd A&. 7 104.98 34.52 49 8827 58 16.9
10th ,, 7 92.85 -12.13 - 12 4975 54 19.1
17th ,, 7 121-78 28.93 30 4894 40 21.5
24th ,, 7 169-53 47.75 38 3464 20 19.8
31st ,, 7 212-72 43.19 25 4807 23 19.0
7th Sept. 7 213.29 0.57 0-3 5204 24 16.1
(-5)
15th ,, 8 202.19 - 11.10
-4 2738 14 17.5
* The fipm in brackets in column 5 give the percentage incrsaes in dry-weight for
the number of day8 stated in column 2.
 G. E. BRIGGS,
F. ' 7 ~ AND
~ ~ C WEST
, 111
Table VIII. -"F'ferdeznhn " Maize grown at Poppelsdorf in 1875.
Total dry- Increase i n Weekly Ratio of Record of
weight of dryweight percentage leaf-area Mean appearance
Date of 1:rowth a sitigle since last increase in Leaf- to d y tempe-a- of 6 3nd 0
harvest ]wio,i plant harvest drp~weight area weig t ture flowers
since last R q . cni. Sq. cm.
1875 Days Gm. Gm. harvest per plant pergm. "C.
11th May 0.294
( - 3.4)*
1st June 21 0.286 - 0.010 -1.1 36.2 126 14.0
8th ,, 7 0.517 0.232 80 77.9 150 19.3
15th ,, 7 1.0'23 0.506 38 177 174 17.1
(74)
23rd ,, 8 1.781 0.758 65 335 188 16.6
30th ,, 7 3.826 2.045 115 730 190 17.0
7th July 7 9.064 5.438 135 1686 187 19.9
(91)
13th ,, 6 17.292 8,228 106 2578 149 18.1
(72)
21st ,, 8 29.704 12.41 63 3984 134 18.8
(85)
27th ,, 54.998 25.29 100 6274 114 17.6
3rd Aug. 73.949 18.95 32 6622 90 16.9
10th ,, 108.68 34-73 46 8453 77 19.1 $ . a n d ?
17th ,, 153.61 45.93 41 8823 57 21.5 flowers
24th ,, 173.18 18.57 12 8258 48 19.8
31sl. ,, 210.37 37.19 21 7090 34 19.0
( - 16.5)
7th Sept. 7 245.09 34.72 - 14.5 9200 38 16.1
* The figures in brocke I i I column 5 give the percentage increase in dry-weight fo-
the number of days stated in column 9.

In fortning a clear picture of .he growth of the plant as presented by

its increase in dry-weight, it is as well to keep in mind the fact thah from
80 % to 30 yo of the dry-weight is the result of the process known as
carbon-assimilationand that the actual percentage of the d y-weight of the
plant derivec from the m'neral constituents of the soil is relatively small
(cf. Hornberger(C), Monnier (171, Rabinovitch(z0) and others(26, 4, 8 & 3))l.
1 Jonrs and Huston(8) give he following figures for the sh of maize at different
periods :
Ash
Date of sampling (percentage of the total dry-
(week from s o w ~ n g ) weight of the plant)
3rd 2.0
Yth 12.''
lth 8.7
14th 6.0
16th 5.3
lSth 4.8
19th 4. 1
20th 4.1
 Fig. 3. Helative growth-rate curves for “Badischar Friih” maize in four successive years.

5
a
.-c
114 Q ~ ~ aAnalysis
~ ~ of~ Plant
~ ~Growth
u ~ i ~ ~

It follows that the relative rate of growth a t any time is almost the
same as the difference between the rates of assimilation and respira-
tion per 100 g. dry-weight a t that time.
We will now proceed to consider the curves. I n following the curves
in Fig. 3 from the date of sowing, there is seen t o be an initial phase
lasting for about three weeks during which the rate of growth is negative,
in other words, the plant is actually losing in weight1. This phase
of negative growth persists until a point in the development of the
plant is reached a t which approximately four leaves have appeared.
During the time occupied by germination, before the appearance of these
leaves, the negative rate of growth is clearly to be attributed to a loss
of carbohydrate through respiration. The order of magnitude of the
loss in dry-weight through respiration in germinating seeds is 3 yoto 6 %
of their dry-weight per day at 16" C. (Garreau(5)). I n the latter part of
the period, where, despite the fact that the plant possesses from 1-4 leaves,
the negative rate of growth persists, it is obvious that any increase in
dry-weight due to assimilation is more than counter-balanced by a loss
in weight through respiration. Evidence obtained by an analysis of
Kreusler's data as to whether the leaves a t this stage perform their
normal assimilatory function, or not, will be considered shortly. After
this initial phase there ensues a short period varying from 1-4 weeks
during which the rate of increase in dry-weight rises rapidly to its
maximum value, followed by a long period constituting the remainder,
and larger part, of the life-cycle of the plant, throughout which the rate
of growth falls off more or less continuously. This falling part of the
curve, however, shows subsidiary maxima.
The question arises of what kind of change in the plant this perfectly
definite type of curve in the main period of growth is an expression. It
is clear that this main rise and fall must be due to an increasing difference
between the rate of assimilation and the rate of respiration per unit dry-
weight in the first phase and to a decreasing differencein the second phase.
The order of magnitude of respiration in terms of dry matter consumed
per week during the main growth period of the plant is probably not
greater than 20 %-40 %, which is the order of magnitude of the loss
in dry-weight through respiration during germination. As against this
the actual percentage increase in dry-weight per week varies from 0 yo
to over 200 %, this being the balance when loss due to respiration is
In the year 1875 the first dry-weight measurement wa8 not taken until the end of
the third week. The average plotted in Fig. 3 gives no indication of the variations in the
individual weekly growth-rates.
 F. KIDD,AND C. WEST.
G. E. BRIGIGS, 115
subtracted from gain due to assimilation, etc. Consequently it is obvioup
that changes in the rate of respiration per unit dry-weight of sufficient
magnitude to affect the rate of growth to the extent observed are in-
conceivable. We must turn therefore to the changes in the rate of
assimilation per unit dry-weight in order to account for the rntLiin rise
and fall which characterises the growth-rate curve. Brief consideration
will show that the rate of assimilation per unit dry-weight is most
probably a function of the amount of leaf-area per unit dry-weight
mainly, and it is interesting to enquire t,herefore to what extent changes
in leaf-area per unit dry-weight correspond with those in the rate of
growth. The values of the ratio of leaf-area to dry-weight throughout
the life-cycle of the plant can be calculated from Kreusler’s data, and
when these values are plotted against time there appears a striking
similarity between this curve and the growth-rate curve (see Figs.
4, 5 , 6 and 7).
From this we may conclude, therefore, that the main rise and fall
shown by the growth-rate curve is merely an expression of the rise and
fall in the ratio of leaf-area to dry-weight.
To return to the question of the assimilation of the young leaves on
their first appearance, an inspection of Figs. 4, 5, 6 and 7 will show that
at this stage the ratio of the ordinate of the growth-rate to that of the
leaf-area curve (which ratio is really a measure of the incrdase in dry-
weight per unit leaf-area)is a negative or very small quantity compared
with the ratio during the main period of high relative rate of growth.
This fact strongly suggests that the assimilatory power of the young
leaves for some time after their first appearance is negligibly small. It
is interesting to find that this inference which is drawn from an analysis
of plant growth, as presented in this paper, is corroborated by direct .
experimentation on the assimilatory power of young leaves (Irving(7)
and Briggs ( 2 ) ) l .
Another point of interest which arises from a comparison of the
leaf-area ratio with the growth-rate curve is that, while the growth-
rate curve exhibits one or more subsidiary maxima in the falling phase,
the leaf-area ratio curve on the other hand falls uninterruptedly.
With regard to these subsidiary maxima exhibited by the growth-
rate curve there is a significant correlation between the times of their
occurrence and the recorded times of the first appearance of the male
and female flowers (see Figs. 4-8). Results obtained with maize by
Morgen (18) and Osswald (19) who worked in conjunction with Kreusler,
1 Also unpublished results for maize.
 Weekly percentage increment in dry-weight
6
Weekly percentage increment in dry-weight

4
2:
Y
E!
a.
(D
c
m
01
4

. i;'
Ratio of leaf-area to dry-weight. Sq. crns. per gm.
0 0
!i E1
2 Ratio of leaf-area to dry-weight. Sq. crns. per g m
 W Weekly percentage increment in dry-weight
a
F Weekly percentage increment in dry-weight

0
Ratio of leaf-ares to dry-weight. Sq. cms. per gm.
3 2 Ratio of leaf-area to dry-weight. Sq. cms. per gm
11s Quantitative Analysis of Plant Growth
are given in Fig. 8. These results are for tops only, not for the entire
plant, including roots, as in the other cases.
It is striking that when there is only one prominent subsidiary
maximum the male and female flowers appear together. These subsidiary
maxima cannot be correlated with recorded variations in any climatic
conditions and consequently it seems safe to conclude that they must
be due t o internal changes.
I n endeavouring to explain these maxima and their correlation with
the appearance of the male and female flowers in terms of assimilation
and respiration there are two alternatives. The first is to suppose that
a t the recorded time of the appearance of the flowers there is a temporary
increase in assimilation per unit leaf-area or a decrease in respiration
per unit dry-weight, or a temporary increase in salt absorption by the
roots. The other alternative is to suppose that during the early stages
of flower development, prior to the first :ecord, the reverse conditions
obtain, in other words, that the minima immediately preceding the
record of the appearance of flowers is to be attributed to ihese reverse
conditions. Since it is a well-known fact t h a t flower developmmt is
accompanied by an increased respiratory activity and also since we have
no evidence that there is an alteration in assimilation per unit leaf-area
connected with flower-formation, the safest conclusion a t present seems
to be that the minima are to be correlated with increased respiratory
activity a t these periods.
Plants grown a t the same time under similar conditions show a
coincidence of the maxima (Fig. 2), but when we compare plants grown
a t different times and under different conditions the incidence of the
maxima varies (Fig. 3). It appears likely therefore that the incidence
of the maxima depends upon external conditions. As attempts to correlate
the maxima with the environmental conditions obtaining at the time of
their incidence were unsuccessful, we have concluded that most probably
the time of the incidence of the maxima is determined by environmental
conditions obtaining a t previous stages in the plant’s development.
Having now considered the whole of the growth-rate curve for maize
i t appears on the basis of the data available that the general form of
the curve and the occurrence of its various maxima are controlled by
internal changes intercorrelated with morphological developments. The
points in morphological development which appear to be significant are
( 1 ) the rise to a maximum and the subsequent fall in the leaf-area dry-
weight ratio, (2) the development of the male flowers, and (3)the develop-
ment of the female flowers. Environmental conditions may influence
G. E. HRIGGS,F. KIDD,AND C. WEST 119

m
120 &?cantitative Analysis qf Plant Growth,
the time relation of these points and thus the time relations of the maxima
on the curve. I n extreme cases the environmental factors may so far
affect morphological differentiation as to cause coincidence of the
maxima.
External conditions, in addition to causing modification in this way
in the general form of the growth-rate curve, must directly affect the
absolute value of the growth-rate, but an analysis of these curves and
attempts to correlate still smaller fluctuations in these curves from year
to year with external conditions have not yielded any definite results.
We shall return to the subject of the effect of external conditions when
dealing later with another form of expressing growth-rate, namely in-
crease in dry-weight per unit leaf-area per unit time.
I n a future chapter we propose to compare the relative growth-rate
curves of other annual plants with those for maize which have been
dealt with above.
AVERAGE
GROWTH-RATE.
A full consideration of all the data presented here will show the
extraordinary difficulty of finding any valid basis for comparing plants
such as maize by means of th& average growth-rate whether the
average is taken over the whole life-cycle, which is of varying length,
or whether arbitrary periods of shorter duration are taken. It is par-
ticularly misleading to compare the average growth-rate for one period
of one plant with a different period of another plant. For example, a
comparison of any two plants by means of their average growth-rate
over a period such as six weeks would be favourable to one, whereas
a comparison over say 12 weeks might be favourable to the other.
In a subsequent chapter dealing with the question of growth-rate in
relation to yield, this point will receive detailed consideration.

SUMMARY.
The series of articles of which this is the first instalment, constitutes
an attempt to formulate methods for the quantitative analysis of plant
growth and to apply these methods to data which have been lying
dormant in the literature for 40 years.
I n the present chapter the relative growth-rate curve, which is the
weekly percentage increase in dry-weight plotted against time, and also
the leaf-area ratio curve, that is, the leaf-area in sq. ems. per g. plotted
against time, have been employed. And as a typical example of an
 G. E. HRICTGP,F. KTDD,AND C. WEST 121
annual plant maize has been selected since data are given by Kreusler
for this plant grown in four successive years.
The first noteworthy resnlt of this analysis is the demonstration of
the fact that the growth-rat,e varies greatly in magnitude a t different
periods in the life-cycle of a plant such as maize in a perfectly definite
manner.
Fig. 9 gives the generalised form of the growth-rate curve for maize
throughout its life-cycle. Although the broad form is that of a Sach’s
grand period curve, it must be noted that i t is not a grand period curve,
since the grand period curve as defined by Sachs is the curve of the
actual increment per unit of time plotted against time and not of
relative increment, that is, increment per unit of matter per unit of time
plotted against, time. On the broad form of the relative growth-rate curve

Fig. 9. Generulised form of the growth-rate curve for mrtize.

for maize are superposed three secondary features, an initial fall, and
two subsidiary maxima on the descending limb.
I n this generalised curve the initial period A-B is the period before
the assimilatory organs are able to counterbalance the lose in dry-weight
due to respiration, and the rate of growth is consequently negative or
nil. The phase B-C corresponds to a phase in morphological develop-
ment during which the leaf-area per unit dry-weight increases to a
maximum. The phase C-F covers the remainder of the life-cycle of the
plant during which the leaf-area per unit drv-weight is continuously
decreasing. The subsidiary maxima D and E coincide with the time of
the record of the appearance of the male and female flowers respectively.
The minima X, Y which precede these maxima, correspond with the
earliest stages of flower development, and are possibly due to increased
respiration during that period.
122 Quantitative Analysis of Plant Growth
The incidence of the maxima is controlled by environmental condi-
tions-not by the environmental conditions operating a t the time, but
by those obtaining at some previous stage in the life-history of the
plant.
The fact that the curve for leaf-area per unit dry-weight throughout
the season (which has been calculated) shows a correspondence with
the growth-rate curve indicates that the physiological basis for increased
and decreased relative rate of growth is a corresponding change in the
assimilating area per unit dry-weight. This point will be dealt with in
the next chapter.
Evidence from -the quantitative analysis of plant growth for maize
indwates that the seedling leaves do not perform their normal assimila-
tory function till some time after their appearance.

(To be continued.)

LITERATURE CITED.
( 1 ) RLAFKMAN, V. H. The Compound Interest Law and Plant Growth. Ann. Bot.
XXXIII, 1919, p. 353.
(2) BRIGGS,G. E. The Development of Photosynthetic Artivityduiing Germination.
Proc. Roy. Soc. (Lond.),B, XCI, 1920, p. 249.
(3) BURD,J. S. Rate of Absorption of Soil Constituents at Successive Stages of
Plant Growth. Journ. tlgrzc. Research, XVIII,2, 1919, p. 51.
(4) DELEANO,M. Etude sur le R81e ef lo Fonction des Sels Mindraux dam la V i e de
la Plante, Geneva. 1907.
(5) GARREAU. Uc la Respiration chez ICY Plantes (1). Ann. Sci. Nut. (Bot.),3e SBr.,
t. xv, 1851, p. 5.
(6) HORNBERGER, R. Untersuchungen uber Gehalt und Zunahme von Sinapis alba
an Trockensubstanz und chemischen Bestandtheilen in 7-tagigen Vege-
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(7) IRVING,A. A. The Beginning of Photosynthesis and the nevclopmcnt of
Chlorophyll. Ann. Hot. SYIV,1910, p. 805.
(8) .TONES,W. J. and HLTSTON, H. A. Composition of Maize at Variouq Stages of
its Growth. Purdue Unic. Aqric. Exp. Sta. Bd1. 175, XVII, 1914.
(9) KIDD, F. and WEST, C. Physiological Pre-Determination. IV. Review of
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(10) KREUSLER, U., PREHN,A. and BECKER,G. Beobarhtungen uber das Wachs-
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Wachsthum der Maispflanze. LuruZw. Jahrh. VII, 1878, p. 536.
(13) __ Lanrlw. J a h b . wir, 1879, p. 617.
(14) ~~TTSCHERLICT:, A. Das Gesetz dcs Pflanzenwachsbums. Landw. Jnhrb. 1,111,
Heft 2, 1919, p. *67.
(15) - Ein Reitraq rum Gesetz des Pflanzenwachstums. Fuhling’s I;adw. Ztg.
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(16) __ Zum Clesetz des Pflanzenwachstums. F,iihling’s LanrZw. Ztg. LXVIII,
Hrft 21/22, 1919, p. 419.
(17) !\~OXNIER, A. Les Mati?res Min,h.ales, et la I d d’Accroiaaemeal des Vdge‘taux,
Grnrva, 1915.
(18) ~ I O ~ W EA. N , Bericht :ibw die im Jahre 1878 an dar Versuchs-station zu Halle
a,. 8. ausgefuhrten Bestimmungrn dcr Trockensubstanz-Zuna~lm~ h i der
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(19) OSSWALD, W. T. Rericht iiber die in1 Jahre 1875 a n der Versuchs-station zu
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Landw. .Jahrh. VIII, 1879, p . 656.
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Rnphanus sntious, Geneva, 1914.
(21) REED, H. S. and H O L L ~ DR., H. The Growth Rate of an Annual Plant
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(22) R,IPPEL,A. Die Wachst.umskurve. Ber. d. Deutsch. Rot. Gesellsch. XXYVII,
Heft 3, 1919, p. 169.
(23) - Die Wachstumskurve der Pflanzen und ihre M a t h e m a h h e Behandlung
(lurch Robertson und Mitscherlich. Fuhling’s L a d w . Ztg. LXVIII, Heft
11/12, 1919, p. 201.
(21) ROBERTSON, T. B. Further Remarks on the Normal Rate of Growth of an
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(25)- On the Natnre of the Aiit,bcata.lyst of Growth. Archiv j. Enturiekelungs.
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