New Phytologist - May 1976 - WALTON - SOME LIMITATIONS ON PLANT GROWTH AND DEVELOPMENT IN TUNDRA REGIONS AN INVESTIGATION PDF

New Phytol.
SOME LIMITATIONS ON PLANT GROWTH AND

DEVELOPMENT IN TUNDRA REGIONS—AN
INVESTIGATION USING PHYTOMETERS
BY D . W . H . WALTON AND R. L L . SMITH
Life Sciences Division, British Antarctic Survey, Monks Wood Experimental Station,
Abbots Ripton, Huntingdon, Cambridgeshire*
{Received 6 October lgj^)
SUMMARY
The use of oats as phytometers for seasonal studies of microclimatic favourability and nutrient
limitations is reported. The effect of age and development on growth parameters in long-term
experiments is illustrated by comparison with data from related short-term experiments.
Regression analyses were carried out on seasonal growth data to discover patterns of growth
and development. Intersite differences were more clearly demonstrated by plants grown in
nutrient-rich media than by those grown in soil and were most marked under poor weather
conditions. Highly signiffcant differences in growth existed between seasons for the nutrient-
treated vermiculite experiments and between treatments for the soil and vermiculite experi-
ments. The use of regression analyses to explain dry matter accumulatioti is discussed, and the
usefulness of phytometers in growth assessment is considered.
; -:..,. INTRODUCTION •, Ai . ,,
In an earlier paper Smith and Walton (1975) discussed the use of three temperate erop
species (oat, radish and turnip) to assess microclimatic limitations to growth in a sub-
polar cHmate. The results were derived from io-day test periods at different times during
the growing season on the subantaretic island of South Georgia (lat. 54°-55°S., long.
36°-38°W.). A serious limitation in this type of experiment is the necessity, for purposes
of comparability, of using plants all at the same stage in ontogenetic development. No
information can therefore be gained about the interactions between development, age
and microenvironment. Experiments were therefore carried out on South Georgia
during the summers of 1969-70 and 1970-71 to investigate the effects of age on growth
under different local climates and, in view of Warren Wilson's (1966) evidence of nutrient
limitations in polar soils, to assess the difference in growth between plants grown in a
nutrient-treated medium and those grown in local soil. ;
MATERIALS AND METHODS

The three sites used for the experiments (Primary, Festuca-heath and Fellfield) were
those referred to by Smith and Walton (1975) and described by Smith (1971). Pre-
germinated seeds of oats (Avena sativa L. cv. Astor) were grown at each site in pots of
vermiculite watered every 2 days with modified Hoagland's solution (Lewis and Greene,
1970), and in shallow drills in untreated grassland soils which resembled Arctic brown
soil as described by Tedrow and Hill (1955).
•Present address: Life Sciences Division, British Antarctic Survey, Madingley Road, Cambridge CB3
oET.
SOI
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5O2 D. W. H. W A L T O N AND R . I. L. SMITH
The soil experiments were carried out only at the Primary and Festuca-heath sites,
soil being almost absent at the Fellfield site. The first of the fortnightly frequent small
harvests (Hughes and Freeman, 1967) was taken when the plants reached the 3-leaf
stage. At each harvest pots were selected randomly from each site to provide fourteen
healthy plants. In each soil treatment the drills were divided into blocks and one,
containing approximately fifty plants, was randomly selected at each harvest. The thirty
largest plants in terms of fresh weight were used for analysis. For each plant from both
treatments, leaf number, tiller number and leaf area were recorded and the dry weight
of leaves, shoots and roots was determined to the nearest 0.1 mg after drying for 3 days
at 8o°C. Although the planting date was the same in each year for both treatments
the date of the first harvest difiêred since this depended on the rate of development of
the plants. Unless otherwise stated all growth analyses are therefore calculated from the
standard baseline of the 3-leaf stage of development and do not take into account the
time necessary for the plant to grow to this stage.
Mean dry weight and total leaf area values were tranformed into natural logarithms to
render their variability more homogeneous with time (Hughes and Freeman, 1967).
Polynomial regression curves were fitted to the transformed data in a stepwise manner
by the least squares method. The significance of the reduction in total variance made
by linear, quadratic, cubic and quartic terms was tested by E values calculated from an
analysis of variance. The ratio of the regression mean square for eaeh polynomial against
the residual mean square was tested by Fisher's F-test. The highest term significant at
the 5% level of probability was normally accepted and the relevant equation used to
describe the increase in log^ dry weight and log^ leaf area with time. At intervals of 5
days fitted values of log^ dry weight and log^ leaf area were substituted in the growth
formulae (Radford, 1967) and values of relative growth rate (i?w). net assimilation
rate {E/^ and leaf area ratio {Fp) were calculated. The curve shape for growth parameters
presented in this paper is dependent upon the regression term originally fitted to the
transformed dry weight and leaf area data. Least significant differences between means,
at the 0.1%, 1% and 5% levels of probability, were calculated for the data sets used in
comparison. A detailed account of this method is contained in Callaghan (1974). A
similar approach to data analysis has been examined by Nicholls and Calder (1973)
and described by Hunt and Parsons (1974).
-:• -, . , : - v y • ' • ' ' ' < ; . • ' • ^ ' • ' t ' O . i f j « i t ' " ' . . > • • y ' K U - - ' s : - ^ i., 1.-'". i \ ' ^ ••'••'' •• •'''^'- ' " • ' - : • ( • > • : ' • . ' . ;.-
RESULTS '^.'-'rt-^v , ,'..''.',,..^^.'".:V'.',..^,.'

Comparison of growth and development
Mean data for morphological measurements and dry weight determinations after a
growing period of n o days in eaeh season are given in Table i, while mean monthly
climatic data, reeorded near sea level close to the field sites, are provided in Table 2.
Two general points are immediately apparent. Firstly, there was a considerable difiêrenee
between the performance of the plants grown in vermieulite and those grown in soil
in the 1970-71 season. This was not apparent in the previous season. Seeondly, there was
a marked difference between sites for most parameters. In both seasons the mean number
of leaves per plant was mueh lower in the soil treatment, although they were larger and
thicker than those of the plants grown in vermieulite. None of the plants grown in
vermieulite produeed maeroscopieally visible inflorescences whilst most of those grown
in soil had immature ensheathed spikes towards the end of the season. Tiller production
appeared to be negatively correlated with the development of infiorescences and, in
iO?
Growth and development in tundra regions 503
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5o6 D. W. H. WALTON AND R . I. L. SMITH
the plants grown in soil, it was very low in comparison with plants in vermiculite. The
above:below ground dry weight ratios reflect major differences between treatments in
dry matter partitioning by the plants. In both seasons the ratios were higher for the
plants grown in soil than for those grown in vermiculite in which root development was
often considerable. Thus, despite the much lower number of tillers and leaves per plant
in the soil treatment, the development of the inflorescence in these plants ensured that a
greater proportion of the dry matter remained above ground than occurred in the plants
in vermiculite. The relatively high nutrient status of the vermiculite appeared to stimu-
late vegetative growth at the expense of reproductive development, while in the soil,
the low mineral status produced the reverse effect.
A difference in plant response to microclimate at the three sites is clearly distinguish-
able in Table i for the first season of the vermiculite treatment, yet in the second season
there was little difference between the sheltered Primary and the exposed Fellfield
sites. On the other hand there was a marked difference in performance of the soil-grown
plants at the Primary and Festuca-heath sites for the latter season. Since there was little
difference in growth in vermiculite between sites in the 1970-71 season, suggesting
comparatively similar local climates, the variation in growth in soil is probably attri-
butable to differences in nutrient status. A chemical analysis of the soils (Table 3)
indicates considerable differences in organic matter, moisture content and N, although
Na, K, Ca, Mg and P levels were similar.
The results obtained in the vermiculite treatment for the two seasons suggest strongly
that the 1970-71 summer was much more favourable for growth than the previous
season. This agrees with the meteorological data presented in Table 2. Mean air tem-
perature was considerably higher during January, February and March in the second
season, and there was a much lower incidence of days with air frost or snowfall. The
climatic data therefore support the hypothesis that phytometers grown in nutrient-
enriched vermiculite can provide a good assessment of the favourability of growing
seasons for comparative purposes. This is, however, not true for the plants grown in
native soil where dry matter production at the Primary site in the two seasons was
similar despite differences in the number and thickness of leaves and in dry matter
partitioning. It would appear that only under high nutrient conditions can an increase
in climatic favourability enhance dry matter production, the growth of the plants in
soil being too severely restrained by edaphic factors for minor fluctuations in climate
to have any significant influence.
Comparison of regressed growth data

While there are obvious inter-site differences for many of the raw data in Table i,
the corresponding differences for the calculated growth parameters are not so clear or so
easily interpreted. Fig. i illustrates the 7?^ patterns for total plant weight at the three
sites in 1969-70. Significant differences between sites occurred only towards the end of
the growing season, the pattern of dry weight production being similar at all three sites.
In 1970-71, as might be expected from data in Table i, growth rates were much higher
than in the previous season.
The regressions used for JP^ (Fig- 2) did not show significant differences between
sites, but those for E'A (Fig. 3) showed large and highly significant differences for most
of the growing period. The seasonal patterns for fi^ show an overall decline with in-
creasing age and deteriorating weather, and support the inter-site differences indicated
in Table i. The patterns for Fj^ are not as expected. The selection of a constant F^
0.2 III
1 ~-—1
0 20 30 40 50
10 60^ ~~~~~-^^
-O.I Days from first harvesf
Fig. I. Regressions of relative growth rate (i?w) of total plant dry weight against time. Oats
grown in vermiculite at three sites, 1969-70. (• • •, Primary; , Fellfield; , Festuca-
heath). Bars indicate least significant differences for, from left to right, P = o.ooi, P = o.oi,
P = 0.05.
l.2r
1.0
0.8
I 0.6
0.4
0.2
20 30 40 50 60 70 80
Days from first fiarvesf
Fig. 2. Regression of leaf area ratio {F/^ against time. Oats grown in vermiculite, 1969—70.
(. . ., Primary; , Fellfield; , Festuca-heath). Bars indicate least significant differ-
ences for, from left to right, P = o.ooi; P = o.oi and P = 0.05.
for the plants at the Primary site appears biologically unrealistic, whilst the rise in F^
at the other two sites towards the end of the season is certainly at variance with normal
growth patterns. Statistically the computer programtne showed that for these two sites
a cubic relationship was the best fit for log^ dry weight and leaf area and this resulted
in the F^ curves shown in Fig. 2. Examination of the original data suggests that this
anomaly may have resulted from an increase in leaf damage at the more exposed Festuca-
heath and Fellfield sites towards the end of the summer.
0.5r-
10 30 40 50
Doys from first harvest
Fig. 3. Regression of net assimilation rate {E/^ against time. Oats grown in vermiculite,
1969-70. (• • •, Primary; , Fellfield; , Festuca-heath). Bars indicate least significant
differences for, from left to right, P = o.ooi, P = o.oi and P = 0.05.
5o8 D. W. H. WALTON AND R . I. L. SMITH
0.6r-
I I I -•••
0.4 -
—-
E 0.2-
1 1 1 1
y 10 20 30 40 50 60 70 80
Days from tirst harvest
-0.21-
Fig. 4. Regression of relative growth rate (i?w) of shoot dry weight against time. Oats grown
at the Primary site, 1970-71. ( , vermieulite; , soil). Bars indicate least significant
differences for, from left to right, P = o.ooi, P = o.oi and P = 0.05.
Analysis of individual plant fractions can sometimes offer quantification of de-

velopmental observations. Fig. 4 compares the growth of shoots at the Pi'imary site in
both vermieulite and soil treatments. The R.^ values for roots and leaves of the plants in
soil became negative towards the end of the season at the Primary site in 1970-71,
whilst values for the shoot remained positive. 7?^ values for all parts of the plants in
vermieulite were positive throughout the season. The production of inflorescences was
observed only in the soil treatment and it is therefore assumed that the negative rates for
roots and leaves were due to translocation of all available material into the developing
infiorescence.
The computer programme fitted cubic and quartic regressions to a considerable pro-
portion of the transformed data on good statistical grounds, and often this resulted in
seasonal patterns that were difficult to explain biologically. Examination of seasonal
trends has usually shown that the quadratic regressions can be more easily explained in
biological terms than can cubic and quartic, although in some cases, it is realized that this
may lead to oversimplification of the plant response particularly under field conditions.
Polynomial curve fitting on purely statistical grounds can lead, through an infiexibility
of application, to completely misleading conclusions.
DISCUSSION
The data presented in this paper and by Smitb and Walton (1975) were obtained for
three specific objectives. Firstly, to investigate the use of phytometers in the quantifica-
tion of the favourability for growth of various habitats with different microclimatic
regimes. Secondly, to assess tbe pattern of seasonal change in growth parameters resulting
from plant age and development, and thirdly, to compare growth in native tundra soils
with tbat in a nutrient-ricb medium.
The use of a temperate cereal species to assess growth in a subantarctic environment
has several important advantages over native species. The native fiora of South Georgia
consists of rather slow-growing perennials which are adapted to long-term survival rather
than rapid annual growth. Oat is a relatively fast-growing annual even under tbe climatic
conditions prevailing on this island. Several assessments of growth can therefore be made
at a standardized stage of development within each season, with the benefit of large
differences in dry weight between harvests if the experiment continues for most of the
growing season. In terms of genetic variability greater uniformity of material is likely to
be obtained from a crop species than from seed collected from native species. This
should permit a smaller sample size at harvests. An important criterion in studies of
productivity and yield is the ease with which the plants can be harvested and processed.
In this respect oats proved very satisfactory, although other plants have been used
successfully (Bonde et al, 1973; Smith and Walton, 1975). Finally, as a technique for
habitat assessment this type of experiment can be used with minimal destruction of the
local vegetation. This feature is of particular importance in polar and alpine regions
where regeneration of native communities is often very slow.
The importance of specifying the ontogenetic stage of the plants is indicated by the
considerable difference between the 7?^ values in three separate experiments at the
same time and at the same site (Table 4). The difference between short-term and long-
term vermiculite-grown plants is attributable to age, and that between the vermiculite
long-term and the soil long-term plants to edaphic limitations. These data underline the
importance of standardization of both developmental stage and growth medium in any
comparisons.
Table 4. A comparison of relative growth rates (^w) of oats
grown in three treatments at the Primary site in 1970-71
,' Vermiculite Vermiculite Soil
(short-term*) (long-term) (long-term)
7-17 Jan. 1971 0.53 0.34 o.zg
20 Feb.-2 March 1971 0.45 0.24 0.19
, * Data from Smith and Walton (1975).
The reasons for using regression analyses in growth studies are succinctly stated by
NichoUs and Calder (1973) and Flunt and Parsons (1974). In their examination of the
application of this technique to data for Poa annua L., Lolium temulentum L., and Atriplex
spp. Nicholls and Calder (1973) found little evidence to support the use of cubic rela-
tionships although the curve-fitting procedure used in the present paper has shown both
cubic and quartic equations to be statistically highly significant in some instances. It is
perhaps relevant that all the sets of data they examined were derived from plants grown
under standardized growth chamber regimes and were not, as in the South Georgia
experiments, subject to the vagaries of the weather. For plants grown under a constant
temperature day/night regime a quadratic relationship may adequately describe the
change in dry weight and leaf area, but it is difficult to see how this is necessarily ap-
plicable to growth and development under field conditions. The South Georgian growing
period (late October to late March) may be interspersed with frequent, short cold spells
with frost and snow whilst the diurnal temperature amplitude is very inconsistent.
The warnings given by Nicholls and Calder (1973) and by Mead (1971) against overfitting
are certainly timely. The use of regression techniques can help to show correlations
between development and production patterns on a seasonal basis but it is not always
easy to combine a satisfactory biological explanation with a statistically significant
regression.
Phytometers are useful in growth experiments where the aims are to assess the climatic
or edaphic limitations on growth rather than the actual production of native species.
Data presented here and by Smith and Walton (1975) have shown that useful information
can be gained from two experimental methods, the results often being complementary.
This technique would seem to offer considerable possibilities for standardized com-
parisons and quantification of plant growth potential at widely separated sites.
5IO D. W. H. W A L T O N AND R . I. L. SMITH
ACKNOWLEDGMENTS
We are indebted to C. Stephenson and J. R. B. Tallowin for assistance in the field and
laboratory, the subdivision of Chemistry and Instrumentation, Merlewood Research
Station, Institute of Terrestrial Ecology, for analysis of soils and to the British Antarctic
Survey, Meteorology Section, for the data presented in Table 2. Our grateful thanks are
also due to Dr T. V. Callaghan who supplied the computer program and commented
on the manuscript, and to Dr J. K. A. Bleasdale and Dr R. Hardwick whose instructive
comments were greatly appreciated. The work was carried out as part of the International
Biological Programme Bipolar Botanical Project. The authors are grateful to the British
Antarctic Survey for permission to publish this paper.
REFERENCES
BONDE, E . K . , FOREMAN, M . F., BABB, T . A., KJELVIK, S., MCKENDRICK, J. D., MITCHELL, W . W . ,
WOODING, F . J., TIESZEN, L . L . & YOUNKIN, W . (1973). Growth and development of three agronomic
species in pots ('phytometers'). In: Proceedings of the Conference on Primary Production and Production
Processes, Tundra Biome, Dublin, Ireland, April 1973 (Ed. by L. C. Bliss and F. E. Wielgolaski), p. 99.
Tundra Biome Steering Committee, Department of Botany, University of Alberta, Edmonton.
CALLAGHAN, T . V. (1974). Ecophysiological studies on bipolar Phleum alpinum L. Arctic Alpine Res., 6, 361.
HUGHES, A. P. & FREEMAN, P. R. (1967). Growth analysis using frequent small harvests. J. appl. Ecoi., 4,
553-
HUNT, R. & PARSONS, I. T. (1974). A computer program for deriving growth-functions in plant growth-
analysis. J. appl. Ecoi., II, 297.
LEWIS, M . C. & GREENE, S. W . (1970). A comparison of plant growth at an Arctic and Antarctic station.
In: Antarctic Ecology, Vol. 2 (Ed. by M. W. Holdgate), p. 838. Academic Press, London.
MEAD, R. (1971). A note on the use and misuse of regression models in ecology, j ' . Ecoi., 59, 215.
NICHOLLS, A. O. & CALDER, D . M . (1973). Comments on the use of regression analysis for the study of
plant growth. New Phytol., 72, 571.
RADFORD, P . J. (1967). Growth analysis formulae—their use and abuse. Crop Sci., 7, 171.
SMITH, R. I. L. (1971). An outline of the Antarctic programme of the Bipolar Botanical Project. In:
Working Meeting on Analyses of Ecosystems, Kevo, Einland, September 1970 (Ed. by O. W. Heal),
p. 51. I.B.P. Tundra Biome Steering Committee and Atlantic Richfield Company, London.
SMITH, R. I. L. & WALTON, D . W . H . (1975). A growth analysis technique for assessing habitat severity in
tundra regions. Ann. Bot., 39, 831.
TEDROW, J. C. F . & HILL, D . E . (195s). Arctic brown soil. Soil Sci., 80, 265.
WARREN WILSON, J. (1966). An analysis of plant growth and its control in Arctic environments. Ann. Bot.,
3°, 383-

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New Phytol.

SOME LIMITATIONS ON PLANT GROWTH AND

{Received 6 October lgj^)

MATERIALS AND METHODS

RESULTS '^.'-'rt-^v , ,'..''.',,..^^.'".:V'.',..^,.'

a w ^ "ft «""« SS*^

I Js-^1 °"d;^ °d"d

vo t^oq -^vo q Tt- N r^ M

Comparison of regressed growth data

Analysis of individual plant fractions can sometimes offer quantification of de-

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