Seedling Establishment

621
SEEDLING ESTABLISHMENT IN VERTICAL GRADIENTS

OF SUNLIGHT
BY J. P. GRIME* AND D. W. JEFFREY
The ConnecticutAgricultural Experiment Station, New Haven,
and Department of Botany, University of Melbourne
I. INTRODUCTION
Although the amount of solar energy intercepted by a community of herbaceous species
may be comparable with that intercepted by forest (Monsi & Saeki 1953), the height of
the shaded stratum will certainly differ. In grassland the shaded stratum is low and is
usually renewed annually by extension of shoots and individual leaves from positions
near the ground. In forests, however, the shaded stratum is high and arises by expansion
of foliage in situ.
Many seedlings grow out of the shaded stratum of grassland within a few days after
germination, while in forests shading persists beyond the seedling phase and may extend
over many years.
Within low vegetation, small differencesin height are associated with large changes in
intensity, direction and quality of radiation, and establishment may depend upon height,
aspect or inclination of the first leaves produced by the seedling. Because vertical
gradients are less pronounced near the forest floor, initial growth in height may be un-
important; here tolerance ratherthan avoidance of shade would appear to have ecological
significance. Hence, radical differences in adaptation to shade may be expected between
woodland and grassland plants.
To recognize mechanisms by which shade is avoided or tolerated we have examined
the reaction of a variety of seedlings to standardized gradients of light intensity.
II. METHODS OF SHADING

The effect of light intensity on growth rate and morphology has been examined by growing
plants under screens of cotton, plastic or metal (Burns 1923; Blackman & Rutter 1948;
Blackman & Wilson 1951a, b; Blackman & Black 1959). In addition to reducinginsola-
tion, the screens modify the temperature of air and root medium, air turbulence and
relative humidity (Waggoner, Pack & Reifsnyder 1959), factors which may themselves
influence growth rate and morphology (Whitehead 1957; Warren Wilson & Wadsworth
1958; Wadsworth 1960).
A further difficultyis the characterizationof the light intensity experiencedby different
species under the same screen. Both the percentage of daylight transmitted through the
screen and percentage of daylight at 'plant height' have been used for this purpose.
Often the term 'percentage daylight' is inadequate. Because of the interception of low
angle illumination by neighbouring plants and screens, gradients in light intensity and
quality are unavoidable (but they can be minimized), and the depth of shade experienced,
and its change with time, will differ according to morphology (Boysen-Jensen 1929).
In particular, important differences in illumination are likely between tall and short
* Presentaddress:Departmentof Botany, Universityof Sheffield.
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622 Seedling establishmentin light gradients
species under the same screen. (For example, seedlings of Castaneamollissimaand Quercus
rubragrowing together were shaded by small screens inside a glasshouse in Connecticut in
November. A paradoxical situation was observed in Castaneamollissimawhere a decrease
in screen fabric transmission from 7-7% to 2.8% daylight increased relative growth
rate from 4-2 to 15-3 mg/g/day. Vertical gradients in light intensity due to shading by
c57,
FIG. 1. Two methods of subjecting seedlings to vertical gradients of light. Above: side
view; below: vertical projection. c = Black cylinder; f = filter of coloured plastic or (for
short-term experiments) leaf discs laminated between plastic sheets; s = supporting stake.
screens and plants occurred beneath the screens, and, in bright sunlight, the intensity
of illumination at the top of the 2-8% daylight (D.L.) screen exceeded that at the
bottom of the 7-7% D.L. screen. At 2-8% D.L., rapid stem extension was induced in
C. mollissima, and foliage was carried up into the brighter light near the top of the
screen. In Quercus rubra no comparable extension occurred, and the slowest growth
rate was measured under the 2-8% D.L. screen.)
The light intensity gradients in this experiment were extreme. Nevertheless smaller
gradients may explain, in part, the conflicting reports of daylight compensation points
to which attention has been drawn by Bourdeau & Laverick (1958).
One approach to the problem of unequal distribution of radiation in shade experiments

J. P. GRIMEAND D. W. JEFFREY 623
is to minimize the gradients by wide spacing of plants (Blackman & Black 1959) or by
supplementary illumination. If photosynthetic efficiency is the subject, experiments may
be carried out on single leaves, the illumination of which may be strictly controlled
(Bohning & Burnside 1956).
Another approach is to standardize the light gradient. Often, as in the present study,
we are in fact concerned with reaction to gradients in light intensity, quality and direction.
Two methods of constructing radiation gradients are illustrated in Fig. 1. The simpler
technique involves growing plants individually in shaded tubes. Each tube is a vertical,
Diameters from top of cylinder Diameters from top of cylinder
6 5 - 4 ---- 3
-------- -------- 2 -- 1-00 0-75 0-50 0-25
" 000
I-T ' ' "15 I i
U
00 Ve ( / 0~~~1
C.~ 5
i2 - G
5Ma 1940 15 25 0st 30 e

20 60 100 120 160
180c-
Vertex anger (v)
to show position of window (w) for given angle p.m., 1
vertex
whole cross-sectional area of cylinders. Measurements carried out at 3.00-3.10
FIG. 2 FIG. 3
FIG. 2. The correlation between vertex angle in a vertical black-walled cylinder and the
incident visible radiation. Radiation measured with flat-window light meter extending over
whole cross-sectional area of cylinders. Measurements carried out at 3.00-3.10 p.m., 1
March 1964, with sky overcast at site of experiment 3 (New Haven, Connecticut) using sec-
tioned tube. Inset: cylinder (c) to show position of window (w) for given vertex angle (v).
FIG. 3. Relationship between vertex angle and hours of exposure to potential sunlight
at centre of a vertical black-walled cylinder. Calculation for 27 July in New Haven,
Connecticut, assuming no obstruction of horizon.
transparent cylinder standing on the root medium and enclosing a single shoot. A
gradient in intensity is created within the tube by means of a layer of black paint on the
inner surface. The intensity and vertical extent of shading depend upon the diameter of
the tube and the height of the painted layer. Control plants are grown without tubes or
in wholly transparent tubes. Because radiation from low angles is excluded from the
shaded portion of the tube, illumination is virtuallyindependentof neighbouringobstacles.
The light collecting propertyof a shade tube may be characterizedby a curvein which the
vertex angle through which incident light reaches the centre of the tube is plotted against
height. In diffuse light vertex angle is linearly related to the quantity of incident radiation
(Fig. 2). On sunny days the relationship does not hold; however, vertex angle is an index
of the potential length of exposure to direct sunlight in parts of the tube to which direct
sunlight is admitted (Fig. 3).
The gradients inside tubes differ in several ways from those in vegetation. Lateral

illumination is largely excluded, and unless the tube is occupied by foliage, spectral
composition changes little with height.
Many of the technical difficulties with shade tubes have been encountered in the use of
small shade screens. They involve reduced ventilation, increased humidity, mechanical
restriction and overheating. In midsummer in Connecticut air temperature in unshaded
tubes exceeded ambient by 12? C and leaves were scorched on contact with transparent
plastic. Overheating in the blackened tube sections was prevented by coating the outside
with white or aluminium paint (Fig. 4).
12
80
70
10
60 - 9-
C50.
:3 E
O I o 7-6/
|0 40I- 4 F6-
"o_ 7 -- )
rn
46
20 1- ' I
/ "
'''
/
/ /I 2
~~~10
1~~- /, 1
'
0 25 30 35 40 1020 30 40 50 60 70 80 9 100
Airtemperature(0C) Full daylight(/o)
FIG. 4 FIG. 5
FIG. 4. Specimencurvesfor air temperaturesat mid-dayin full sun, August, Connecticut,

outside tubes ( ... ), within clear cylinder(-- -) and within shaded tube with outer coat
of aluminiumpaint ( ). Temperaturemeasuredusingthermocouplesat the centreof the
cylinders.
FIG. 5. Gradientsof light intensitywithinglass tubes,unshaded(a): with 6 cm shading(i),
and 12 cm shading(i). Measurementsin diffuselight, made with flat-windowlight meter
extendingover whole cross-sectionalarea of cylinder.
III. EXPERIMENTS
Three groups of species were used:
Experiment (1) five grasses
Experiment (2) five dicotyledons J from grand n Derbyshre, England.
Experiment (3) ten woody species of the Connecticut suburban forest, U.S.A.
Experiments 1 and 2 were carried out at Sheffield and 3 at New Haven.
A. Experiment 1
Design: Five species were subjected to three shade conditions on two soil types in
five randomized blocks.

Species: The grasses were all of common occurrence in Derbyshire, but differed
markedly in quantity of seed reserve, growth form, and habitat (Table 1).
Soils: Two soils from Derbyshire were used: a rendzina (pH at field capacity 7-1)
from the top 10 cm of a stabilized scree in Coombsdale and a highly fertile brown earth
(pH 6-4) from the overburden of a limestone quarry near Buxton.
Shade treatments: Each shade tube was made from a 15-0 x 2-5 cm glass boiling tube
with a 2-3 mm drainage hole punched in the base. The outside received first a black and
then a white coat of paint. Before painting, a strip of black adhesive cloth tape, 2 cm in
width, was applied along the length which was to be painted. The tape served as a shutter
over an observation window. Tubes were painted to heights that provided 0, 6 and 12 cm
shade.
Table 1. Characteristicsof the species in experiment 1

Seed reserve* Form and height of Habitat
Species (mg dry wt) vegetativeshoot in Vegetation Soils
full daylight
Arrhenatherum 2-98 Erect Tall grassland and
elatius 60-120 open screes
Festucarubra 0'54 Suberect Tall and short Calcareous
10-70 grasslandand turf mildly acid
Holcuslanatus 0-37 Erect Tall and short soils
20-60 grassland
Open woodland
H. mollis 0-20 Erect Mediumand short
20-60 grassland
Open woodland
Deschampsia 0-20 Cushion-like Heaths, moors and Acid soils
flexuosa 25-40 open and closed
woodland
* Caryopsisminus testa and ovary wall.
The penetration of light into the tubes was determinedin diffuse light (Fig. 5) by means
of a pair of matched Weston photoelectric cells.
Procedure: The soils were air-dried, mixed thoroughly and sieved to pass 2 mm. The
tubes were filled to 3 cm, the drainage hole first being covered with a disc of plastic cloth.
Seeds of each species were germinated in daylight on filter paper, and on emergence of the
radicle immediately transferred to the tubes. The tubes were supported by 2 cm high
wooden racks which held them erect at a minimum of 4 cm apart in a quincuncial
arrangement.The experimentwas carried out in a glasshouse with temperaturefluctuating
between 15 and 20? C over the period November-December 1961. Natural illumination
was supplemented by warm white fluorescent tubes suspended 4 ft over the bench and
providing a daylength of 18 h. De-ionized water was added to the soils every 2 or 3 days.
At intervals, the seedlings were examined through the observation slit and measure-
ments of their height were made by inserting a mm scale into the tube. The dry weight of
each seedling was determined after 5 weeks.
Results: Of 150 seedlings in the experiment, nine failed to survive transplanting;
these were the only fatalities to occur. Since no significant effects of soil type were
detected, the shoot heights and yields presented are combined from rendzina and brown
earth.

(a) Height
At 10 days, height in full daylight was in the order Arrhenatherumelatius > Holcus
lanatus > Festuca rubra > Holcus mollis > Deschampsiaflexuosa. Shading increased
height in only three of the species, Arrhenatherumelatius, Holcus lanatusand Deschampsia
flexuosa (Fig. 6). At harvest, a reduction in height under 12 cm shade was measured in
Festuca rubra and Deschampsiaflexuosa and observed in both species of Holcus.
Arrhenatherumelatius showed the most rapid rate of height growth and, in comparison
with controls, a marked increase in height on shading. This species has large seeds.
16r
(a) T
12 -----------
10-
o lS _
168
2 /// ^-Y -N.S..
(1
.NS.(S) N.S.
^ aftr
r o
N.S.NN.S.
DayN.S.
Days after germination
FIG. 6. Growth in height in seedlingsof five grasses,in 12 cm clear glass tubes (D); glass
tubes with 6 cm shading(z) and 12 cm shading(*). (a) Arrhenatherum elatus; (b) Festuca
rubra;(c) Holcuslanatus;,(d) H. mollis; (e) Deschampsiaflexuosa.Arrows show occasions
when statisticaltests were madeof the significanceof differencesof shadedtreatmentsfrom
clear N.S. == not
controls. N.S.
glass controls.
clear glass not significant; two and
one, two
significant; one, and three
three dots
dots represent P<0'05,
represent P<0.05,
P << 0.01
P 0-01 and 0 001 respectively.
and PP << 0.001 respectively.
In6 cmshade yields of dry weight decreased by more than 30 (P<0001) in each
Results with other species show, however, that the initial rate of height growth is not
In height
(Table 2). upon
speciesdependent 12weight
the cm seedlingsoffive
treatment,
the yieldFestuca
seed reserve. de12clearrubra
cm glass tubes
);dfurther.glass 054 mg)
always
always
The
dependent upon
reductionubes
the weight of the seed reserve. Festuca rubra (seed
(seed
in yield 6caused by cm shading were inatherselyrelated to initial
weight 0.54
weight mg)
height
elongated slowly in fall in
light and shade, while Deschampsiaflexuosa(0.20
Deschampsiafiexuosa (0a20mg) responded
10 days (Fig.c)
toafter
moderate Holcus lanatus;
shading a (d) flexuosa,
H. mollis;
withDeschampsia
marked e) in hArrowever,
increase rate of height in shade was considerably
yield growth.
higherthanexpestatistical
tests were
mof regression fitting thedifferences
the of shadedtreatmentsfromThis
(b) Dry
(b) Dry weight
weight
In 6 cm -shadeyields of dry weight decreased by more than 30% (P<0.001) in each
species (Table 2). In the 12 cm shade treatment, yield declined further.
The reductions in yield caused by 6 cm shading were inversely related to initial height
growth. Log (yield in 6 cm shade/yield
shade/yield in sun) was linearly related to shoot height
height attained
after 10 days (Fig. 7). In Deschampsiafiexuosa, however, yield in shade was considerably
higher than expected from the regression fitting the data from the other species. This

suggests that D. flexuosa was adapted to shade by some mechanism other than rapid
elongation. D. flexuosa grew slowly in full daylight (Table 2), a feature associated with
shade tolerance by Went (1957).
Table 2. Yield at 5 weeks of seedlings grown in light gradients 0, 6 and 12 cm

in height
Yield
Dry wt (mg) Unshaded(%)
Species 0 cm shade 6 cm shade 12 cm shade
Arrhenatherum elatius 13-8 66 48
Festucarubra 3-6 15 10
Holcus lanatus 5-4 40 12
H. mollis 3-0 12 11
Deschampsiaflexuosa 2-1 34 17
All reductionsin yield on shadingare significantat the 0-001level.
1'8 Arrhenatherum
elatius
1-7
"~~_ en 0-Holcus
cEX * lanatus
U'1
.]15o Deschampsia
^flexuosa
o
o 1-3-
12
1-2 * Festuca rubra
*Holcus mollis
2 3 4 5 6 7 8 9
Heightof seedlings at 10days(cm)
FIG.7. The relation between the percentagereductionin dry weight of seedlings after 5
weeks in 6 cm shadingand their height at 10 days.
B. Experiment2
The design, shade tubes and soils were the same as in experiment 1. In addition to the
total height, the lengths ofhypocotyl, cotyledons, leaf laminae and petioles were measured
regularly. The experiment was carried out in August-October, 1961 and was harvested
after 8 weeks.
The species grown in experiment 2 were all dicotyledons, familiar in Derbyshire
grassland (Table 3).
Results: There were no significant effects of soil type. Data were therefore pooled as in
experiment 1.
(a) Height
Betonica officinalis and Rumex acetosa, which are frequent in dense grassland,
made effective growth in height in the shade treatments. In Betonica officinalis, the tall

habit of the seedling appeared only in shade, while Rumex acetosa elongated rapidly in
both full light and shade (Fig. 8). Despite an initial increase in height growth, shaded
specimens of Arenaria serpyllifolia and Hieracium pilosella failed to exceed 2-0 cm
(Fig. 17). The height of Lotus corniculatus varied within the same shade treatment;
while the majority failed to reach 2-0 cm, three seedlings emerged from the 6 cm shade.
Scale diagrams of the seedlings (Figs. 9-12) show the loci of elongation. There were
two distinct phases in height growth in L. corniculatus, Rumex acetosa and Betonica
officinalis. Initial rapid extension growth took place in the hypocotyl. In the second
phase internodes in Lotus corniculatus,petioles in Rumex acetosa and both internodes
and petioles in Betonica officinalis expanded. In Arenaria serpyllifolia and Hieracium
pilosella effective response in height was confined to the hypocotyl although cotyledon
stalks and petioles extended slightly in the latter (Fig. 12).
Table 3. Characteristicsof the species in experiment2

Seed reserve* Form and height (cm) of Habitat
Species (mg dry wt) vegetativeshoot in full Vegetation Soils
daylight
Lotus 1.67 Mat of prostrateor de- Discontinuousor short Calcareous
corniculatus cumbentshoots grasslandand turf and mildly
3-20 acid soils
Betonica 1.62 Rosette of + ascending
officinalis long or short-stalked
leaves
5-25 Grasslandof medium Mildly acid
Rumex 0.72 Rosette of+ ascending height and open places soils
hi woodland
acetosa longor short-stalked
leaves
5-25
Hieracium 0-15 Stolons bearingrosettes Discontinuousgrass-
pilosella of subsessileleaves land and heavily grazed
1-10 turf Calcareous
Arenaria 0-08 Slenderdecumbentor Bare ground, arable acd silys
serpyllifolia ascendingshoots fields, walls, bare
2-5-25 patches in grassland
* Seed minus testa.
The species capable of sustained height growth in the tubes had comparatively large
seeds. In Lotus corniculatus,however, extension of the stem produced an unstable structure
liable to fall and succumb to soil fungi. Exceptional shoots which emerged from 6 cm
shade were supported by the tube.
Increased petiole length in shade in Betonica officinaliswas correlated with a reduction
in size of the laminae (Fig. 10).
(b) Survival and yield

Shading caused fatalities in all five species (Table 4) and mortality under 12 cm shade
reached 100% in Lotus corniculatus and Arenaria serpyllifolia, two species frequent
in relatively unshaded habitats. Of the three species with survivors in the 12 cm shade
tubes Rumex acetosa, the most productive species in full light, showed the greatest
reduction in yield in 12 cm shade.

6
(a) (b) (c)
5 -l
4 -
I I I I I I
10 20 30 10 20 30 40 50 10
6
(d) (e)
2 N.S.
1 /
Arrowsshow occasionswhen statisticaltests were made of the significanceof differencesof

shaded treatmentsfrom clear glass controls. N.S. = not significant;one, two and three
dots representP< 005, P < 001 and P < 0001 respectively.
weight of seedlings after 8 weeks growth in light gradients,

Table 4. Survival, and dry
60, and 12 cm in
height
Yieldt
No. of fatalities* 0 cm 6 cm 12 cm
Species 0 cm 6 cm 12 cm shade shade shade
shade shade shade (Dry wt (mg)) (% Unshaded)
Lotus corniculatus 0 7 10 10-7 17
Betonica officinalis 0 0 2 10-7 27 13
Rumex acetosa 0 4 6 23-0 13 4
Hieracium pilosella 0 2 8 4-8 12 9
Arenaria serpyllifolia 0 7 10 10-8 3
* Maximumpossible 10.
t Each value tabulatedis the mean for survivingseedlings.

5r
4 (a)
3-
2 -
1 ,\#[. \*.
5r
4- (b)
3-
,-
1-
3, (c)
0 0
0 5 10 15 20
controls, plants were grown both in transparent tubes and in the open. The experiment
was replicated in five randomized blocks.
Species: With the exception of Castanea mollissima, a native of China, the species
used in experiment 3 commonly occur in the second-growth forests and suburbs of New
England. The group includes several species which regenerate in moderate forest shade
(Table 5). In addition to these 'tolerant' species, there were several pioneer species which
do not persist below a closed canopy.
Soil: The soil was a Cheshire (Conn.) fine sandy loam of high fertility.
Shade treatments: The tubes were 55 x 8-5 cm cylinders of transparent plastic. They
were masked to heights of 0, 20, 38 and 55 cm with black paint. Each tube was supported
by a wooden stake.
Procedure:The soil was sieved to pass 1 0 cm and mixed thoroughly. After the addition
of lime and fertilizer, it was placed in 250 containers of 4600 cm3 capacity. The containers
were arranged outdoors with minimum distance of 33 cm between their centres. Seeds
were germinated on Cheshire loam, and when the radicle emerged, a seedling was set in
the centre of each container and a tube placed over it. The height of each seedling was
recorded and dead specimens were replaced by freshly-germinated seedlings at weekly
intervals. Thus as many as twelve fatalities were possible in a single container during the
experiment. Due to shortage of seed, dead seedlings were not replaced in Ailanthus
altissima.

(b)
r, / . i I i.. I.. ..
10 15 20 25 30 35 40 45 50 5 60
5
@
23- (a)
2-/
3
-2v~vVV ?(b)
0
2 (C)
25 30 4050 55
5 10 15 20 35
2r (b)
. I 1i - I -
IT I
-
0
' T.
0 tT, T T T7
20 25
5 10 15
Days after germination
FIGS. 10-12. Mean length of hypocotyl, cotyledons, internodes, petioles and laminae of
seedlings of Betonica officinalis (Fig. 10), Rumex acetosa (Fig. 11) and Hieracium pilosella
(Fig. 12). (a) Unshaded control; (b) 6 cm shade; (c) 12 cm shade.

Despite daily watering, drying of the top cm of soil was unavoidable. Crusting of the
soil surface was prevented by hoeing at 4 and 8 weeks The experiment was performed
in 1963 between 7 June and 29 August. All survivors were harvested after 12 weeks.
Table 5. Characteristicsof the species in experiment 3

Seed reserved* Frequencyof
Species (mg dry wt) Habitat colonized by seedlings
regenerationbeneath
closed canopy
Betulapopulifolia 0.1 Abandonedarableland, open woodland Very low
B. lenta 0-7 Open woodland Low
Rhusglabra 1.5 Abandonedarableland, neglectedpasture Very low
Acer rubrum 7-3 Low densityforest Moderatelyhigh
Liriodendron tulipifera 8-5 Forest clearings Low
Ailanthusaltissima 9-3 Roadsides,wasteplaces,open woodland Very low
Pinus strobus 14-8 Abandonedfields,open woodland Low
Gleditisia triacanthos 40-3 Forest clearings Low
Quercusrubra 1969-2 Moderatelydenseforest High
Castaneamollissima 2015-1 Moderatelydenseforest High
* Seed minus testa.
70
(a) (c) (e)
60 '
-
0
0 ---------- ______________|-------------------------- ------------ ---

50
20 -------,-
0 2 6 8 10 12 0 2 6 8 10 12 0 2 6 8 10 12
- L(b) (d) (f)
.0 60 - -
50 -
40-
30
10 - m
0 2 4 6 8 10 12 0 2 4 6 8 10 12 0 2 4 6 8 10 12
Weeks
FIG. 13. Growth in height in seedlings of six tree species in control (no tube) (0) and in
55 x 8-5 cm plastic cylinders,unshaded(w), and with shade strataof 20 cm (B), 38 cm (1)
and 55 cm (1). (a) Castanea mollissma; (b) Ailanthus altissima; (c) Quercus rubra; (d)
Liriodendron tulipifera; (e) Gleditzia triacanthos; (f) Rhus glabra.
Results: Height of Castaneamollissima, and Quercusrubrawas greater in the unshaded

tube than where there was no tube (Fig. 13), but differences in height and yield between
these two treatmentswere small; thus suppressionin the shade tubes may not be accounted
for by effects of the tubes other than those resulting from shading.

(a) Height
Castanea mollissima was the only species to emerge from 55 and 38 cm shade tubes.
All species showed some height growth in shade (Table 6). With the exception of
Quercusrubra and Betula lenta the height attained in the two most severe shade treat-
ments was linearly related to the logarithm of the mean weight of seed reserve (Fig. 14,
Table 6). The relatively poor height growth of Q. rubra was most apparent when we
compare height growth with that of Castanea mollissima, a species of similar seed weight
(Fig. 13).
Castanea
mollissima
2.5-
E Quercusrubra
E
._
*Gleditsiatriacanthos
20 * Pinus strobus
_.
E
:3 Acer rubrum
xE Liriodendron
tulipifera
2 I - Rhus glabra
*Betula populifolia
Betulalenta
1'0 05 0 0'5 10 1-5 2-0 2-5 3-0 35

Log Mean weight seed reserve (mg)
FIG. 14. Log maximum height attained by seedlings within 55 cm shade stratum plotted
against log mean weight of seed reserve in nine tree species.
(b) Survivaland dry weight

The death-rate of the seedlings varied with species and shade treatment (Table 6).
In all species except C. mollissima fatalities increased with height of shading. In 55 cm
shade, the number of fatalities per container was inversely correlated with the weight of
seed reserve (Fig. 15).
In the two species of Betula, death-ratewas high in the two unshaded control treatments
and was apparently the result of droughting of the surface soil. The downward extension
of roots was initially slow in both of these species. Fatalities were fewer in Betula populi-
folia and Rhus glabra in the unshaded tubes than where tubes were absent.
From a comparison of yields measured on survivors of the first sowing (Table 6) and
the initial weights of seed (Table 5) it is clear that species confined to the bottom of the
55 and 38 cm shade cylinders (i.e. all except Castanea mollissima) were below the com-
pensation point. It may be supposed, therefore, that under these conditions, longevity
was determined by the quantity and rate of mobilization of seed reserve, a view supported
by the correlation between seed weight and death rate (Fig. 15). However, since this same
correlation existed in the 20 cm gradient in which survivors of all species showed a net
gain in dry weight at the end of the experiment (Table 6), we may suspect that there were
J.E. E

Table 6. Survivaland shoot height and weight at harvest of seedlings grown in full sunlig
Fatalitiesper container* Maximumheightattained M

(mm) (meanof container
Species maxima)
No 0 20 38 55 No 0 20 38 55 N
tube cm cm cm cm tube cm cm cm cm tu
Betula populifolia 2-8 5-0 3-2 5-0 4-6 11 12 13 11 18
B. lenta 2-6 1-8 2-8 4-2 5-4 14 11 18 13 13 1
Rhusglabra 0-6 1-6 3-0 4-6 4-8 240 137 58 52 41 2534
Acer rubrum 0-4 0-0 2-4 2-4 3-2 73 128 51 50 54 54
Liriodendron tulipifera 0-6 0-6 1-0 2-0 3-4 58 81 78 43 51 39
Ailanthus altissima - -- -15770t
Pinusstrobus -t 1-2 2-2 3-0 3-6 62 51 73 81 93 1
Gleditsia triacanthos 0-0 0-0 0-2 0-6 1-8 349 408 104 92 133 515
Quercusrubra 0-0 0-0 0-2 0-6 1-4 182 266 239 213 226 629
Castaneamollissima 0-0 0-2 0-2 0-2 0-0 252 380 492 566 592 958
* As seedlingsdied they were replacedexcept in the case of Ailanthusaltissimaand this affectsth
t Indicatesyield measuredon a survivorof the first sowing, i.e. 12 weeks old.
I Exposed seedlingsof Pinus strobuswere decapitatedby house sparrows.

disease factors involved in the shade fatalities and that large-seeded species were resistant
to these or succumbed to them less rapidly.
Numerous observations leave no doubt that soil fungi killed many seedlings. In all
species except C. mollissima, Quercus rubra and Gleditsia triacanthos, damping-off was
observed throughout the experiment in etiolated seedlings in 55, 38 and 20 cm shade.
Betula lenta
* Rhusglabra
Betulapopulifolia
4-
* Pinus strobus
Liriodendron ?
tulipifera
a-^O4 ?Acer rubrum
2
*Gleditsia triacanthos
1 _Quercus rubra *
Castanea
mollissima
I I
, I I*
-0 5055 0 1-0 1-5 2-0 2-5 3-0 35
log Mean wt.seed reserve (mg)
FIG. 15. The relation between death rate (mean number of fatalities per container in 12
weeks) in 55 cm shade stratum and log mean weight of seed reserve in nine tree species.
IV. DISCUSSION
The investigation provides evidence of two distinct responses to shade which may be of
ecological significance. The first is shade avoidance by rapid initial growth in height.
The second is shade tolerance, apparent in the persistence of seedlings in dim light.
A. Shade avoidance by initial height growth

For effective height growth to take place soon after germination, either as a normal
feature of development or as a response to shading, both a source of energy, and extension
sites in the seedling are required.
(a) Source of energy

In all the species examined, growth from 0 to 10 days was largely independent of
shade, indicating that during this period structural materials and respiratory substrates
were derived from the seed. In some of the large-seeded species, e.g. Quercus rubra,
growth was measurable in seedlings which had been below the compensation point,
i.e. in 38 and 55 cm shade tubes for as long as 4 weeks (Fig. 13, Table 6). In several
species with small seeds, e.g. Arenaria serpyllifolia, Betula spp., growth ceased abruptly
after about 10 days in deep shade in spite of the fact that extension sites were available
in the shoot.
In each experiment, greatest elongation in response to shade was observed in species
having large seeds. However, in several species, e.g. Quercus rubra,Lotus corniculatus,
the height attained in shade was considerably lower than that expected if quantity of
seed reserve was the only factor limiting height growth.

When seed reserve is exhausted or unavailable, height growth is dependent upon
photosynthate for energy. If an extension site is available, height growth will then be
determined by two radiation-dependent processes: (1) supply of photosynthate (related
directly to intensity), and (2) inhibition of cell wall expansion (inversely related to
intensity). In many cases we can determine whether rate of height growth is limited by
photosynthate or by photoinhibition of extension by examining the effect of shade on
the rate of height growth. Where photoinhibition is limiting, a decrease in height incre-
ment occurs with increasing light supply, e.g. Castanea mollissima (Fig. 13), Betonica
officinalis (Fig. 8), Rumex acetosa (Fig. 8, 0-10 days). In contrast, height growth is
accelerated with increasing insolation when photosynthate is limiting, e.g. Gleditsia
triacanthos(Fig. 13), Rumex acetosa (Fig. 8, 10-50 days).
(b) Extension sites

(1) Dicotyledons. When the increase of height with shading is due to an effect on cell
expansion, the reaction is limited to certain plastic parts of the shoot. The position and
number of potential sites for extension differ according to the species and their maturity.
In epigeal dicotyledons, extension immediately after germination is limited to the
hypocotyl. Soon plasticity is lost in the hypocotyl and the capacity for height response is
assumed by a succession of sites in the shoot. These consist of internodes or petioles.
Net height response to shade is, therefore, an aggregate that will depend upon (i) number,
(ii) distribution, (iii) sensitivity, (iv) plasticity, and (v) stability of the individual extension
sites.
(i) Number: In experiment 3, a great difference in height response was apparent
between Castanea mollissima and Quercus rubra, which have similar seed weight. In 38
and 55 cm shade, extension occurredin Castaneamollissimain fifteen to twenty internodes
which were produced continuously throughout the experiment. In Quercus rubra, the
shoot apex was dormant for long periods, and in both full daylight and shade only two
or three internodes were developed. Hence, poor height growth in Q. rubrawas due to the
fact that no extension sites were available at certain periods.
(ii) Distribution: A distinction may be drawn between species in which the extension
sites were the internodes, e.g. Castanea mollissima and Arenaria serpyllifolia, and those
in which extension is mainly petiolar, e.g. Rumex acetosa and Betonica officinalis. In
the former, height responses are cumulative. This is not the case, where growth and
extension are confined to the petioles since all leaves originate at the level of the soil.
It is apparent from Fig. 10, however, that both hypocotyl and stem may contribute to
the height growth of shaded seedlings of plants which remain as rosettes in high light
intensity.
(iii) Sensitivity: Rumex acetosa and Betonica officinalis have short petioles when
grown at high light intensities in midsummer. In the autumn sunlight of experiment 2,
petiole extension occurred in unshaded seedlings of Rumex acetosa but not in Betonica
officinalis(Figs. 10 and 11). This indicates a differencebetween two species in the amount
of radiation necessary to suppress extension.
(iv) Plasticity: Even under conditions most favourable for etiolation species differ
greatly in response. For example, in petioles there is a range in plasticity from zero in
Lotus corniculatus(Fig. 17) to a very high value in Betonica officinalis (Fig. 10). Such
differences may arise from disparities in both the number of cells involved and the
plasticity of individual cells.
An example of exceptional plasticity was observed in a recent experiment with Plantago

lanceolata (which in closed grassland survives as a rosette of ascending leaves of up to

40 cm in height). Seedlings established in shade tubes of the dimensions used in experiment
1 displayed remarkable plasticity in the cotyledons and first two leaves. Whereas all the
foliage in control plants in full daylight were ovate-lanceolate, the two youngest leaves
26
24-
22
20-
18-
16-
12
H C L1 L2 L3 H C L1 L2 L3
FIG. 16. Scale drawing of hypocotyl (H), cotyledon (C) and leaves (L) on two representa-
tive seedlings from an experiment in which seedlings of Plantago lanceolata were grown in
2-5 cm diameter clear glass tube, unpainted (left) and painted to 12 cm (right), for 6 weeks.
F S F S F S F S F S
Betonica Arenaria Hieracium Lotus Rumex
officinais serpyllifolia pilosella corniculatus acetosa
FIG. 17. Silhouettes of seedlings of five herbaceous species after 8 weeks growth in full
light (F) and in 6 x 2.5 cm shade tubes (S).
produced in a 6 cm shade tube were linear-lanceolate and extended to double the length
of the control leaves (Fig. 16). The third and fourth leaves of shaded seedlings were ovate-
lanceolate. Experiments are in progress to determine whether the eventual appearance of
broad leaves in the shaded plants is due to an inherent difference in plasticity between

the second and third leaves or is a consequence of penetration of the shade stratum by
preceding leaves.
(v) Stability: Etiolation may produce stems, e.g. in Castanea mollissima, or petioles,
e.g. Rumex acetosa, which are mechanically unstable. In experiment 2, the shoot of
Lotus corniculatuswas liable to collapse after slight extension.
(2) Monocotyledons. In the grasses examined in experiment 1 height growth and
response were apparently controlled by mechanisms similar to those described for the
dicotyledons.
Height response in the large-seeded species, Arrhenatherumelatius, was limited by
photoinhibition over the whole experimental period. This was apparent in the increased
rate of height growth in shaded seedlings (Fig. 6). In contrast, etiolation in the 12 cm
shade tubes was confined to a brief initial period in Holcus mollis and Deschampsia
flexuosa. Etiolation in the latter species was sufficient to cause emergence of foliage into
full daylight before seed reserves were exhausted.
A striking feature of experiment 1 was the absence of a height response to shading in
Festuca rubra. This is remarkable in view of the response, under the same conditions,
of the small-seeded species, Deschampsiaflexuosa.
The effect of shade on the morphogenesis of grasses was not examined in detail. It
would be of particular interest to describe the role of the intercalary meristem in the
shade responses of grass seedlings.
B. Shade tolerance
(a) Resistance to disease
In both experiments with dicotyledons many fatalities occurred and these were most
frequent in shade treatments. Results obtained by Vaartaja (1952, 1953, 1962), Vaartaja
& Cran (1956) and Wardle (1959) have established that soil fungi are important factors
in the elimination of shaded seedlings. In experiment 3, damping-off was observed in
shaded seedlings both above (20 cm shade tube) and below (38 and 55 cm shade tubes)
the compensation point.
Among the tree seedlings examined, longevity in shade was greatest in species having
large seeds (Fig. 15). There is as yet no clear evidence of a mechanism relating resistance
to fungal infection with quantity of seed reserve. In considering the influence of large
seeds on survival in shade, account also must be taken of the greaterinitial height growth
(Fig. 14) and lower relative growth rate of large seeded tree species. This last point is
particularly clear if we compare the yields of Rhus glabra and Ailanthus altissima with
those of two large seeded species, Quercusrubraand Castaneamollissimain the unshaded
treatment of experiment 3 (Table 6).
(b) Morphologicalresponses increasinglight interceptionin shade

Phenotypic adaptations which are induced by shading and which increase the photo-
synthetic efficiency of seedlings in shade are well known (Blackman & Rutter 1948;
Blackman & Wilson 1951; Bordeau & Laverick 1958), and two were observed in
experiment 3. Both increased the energy intercepted by the shoot. In the first, there was
an increased dispersion of chloroplasts in broader, thinner leaves, e.g. C. mollissima
and Quercusrubra.In the second, laminae approached the horizontal by deflection down-
wards in some species, e.g. Gleditsiatriacanthos,and upwards in others, e.g. Acer rubrum.

(c) Metabolic rate

The survival of seedlings with foliage confined to the bottom of the tubes in experi-
ments 1, 2 and 3 was greatest in species which grew slowly in the unshaded treatments.
Shade mortality in species of contrasting growth rates is compared in Table 7. Arenaria
serpyllifolia and Rhus glabra grew very rapidly in full daylight and had poor survival in
shade in comparison to slower growing species. This contrast supports the suggestion
of Went (1957) that the most important adaptation of shade tolerant plants arose by
selection for slow respiration in shade with concomitant selection for slow metabolic
rate, the latter imposing low productivity in sun and shade.
Table 7. Estimated growth rate in full sunlight and mortality in shade

Species Seed weight(mg) Yield in full sunlight Total no. fatalitiesafter 5 weeks
at 5 weeks (mg) in 12 x 2-5 cm shade tube (%)
Arenaria serpyllifolia 0-08 10-83* 100
Betonica officinalis 1-62 10-72 20
EXPERIMENT3
Species Seed weight(mg) Yield in full sunlight Mean no. of fatalitiesper container
at 12 weeks (mg) after 12 weeks
38 x 85 cm shade 55 x 85 cm shade
tube tube
Rhus glabra 1-53 16060-0* 4'6 4-8
Quercus rubra 1969-20 3750.0 0-6 1'4
* When consideredin relationto initial weight of seedling(inferredfrom weight of seed reserve)these
yields indicatevery high relativegrowth rates.
There is some evidence which supports this hypothesis. Bohning & Burnside (1956),
using single attached leaves, reported that photosynthetic rates in tolerant species were
exceeded by those measured in intolerant species, even when measurements were made
at low light intensities. Grime (1965a), using small seedlings in which mutual shading
was slight, found that relative growth rates in full summer sunlight were consistently
slower in shade species, e.g. Acer saccharumand Pachysandra spp. than in sun species,
e.g. Paulownia tomentosa and Helianthus annus. In addition, respiration rates of leaf
discs from plants growing under high light intensities were found to be slower in the
former species.
C. Extrapolation of the experimentalresults to thefield

(a) Shade avoidance
Rapid elongation was observed in shaded seedlings of several grassland species
(Arrhenatherumelatius, Betonica officinalis, Rumex acetosa and Plantago lanceolata).
Since these, unlike many species in experiments 1 and 2, commonly occur in ungrazed
grassland, we suspect that seedling establishment in closed turf is possible in these
species and that rapid emergence from shade permits the establishment.
Professor C. D. Pigott has found that rapid emergence allows colonization of stabilized
screes by Arrhenatherumelatius in Derbyshire, England. Seeds are washed down between
the rocks, and germinate in humid air pockets which may be a considerable distance
below the scree surface. From this position, the first leaf grows upward to the light through
a distance of up to 10 cm.
Since extension sites are available in most seedlings, initial response in the majority of

species is determined by the size of seed reserve (Fig. 13). However, caution is necessary
for there is evidence to support the contention of Shirley (1943) that large seed reserves
are used for the initial development of the root rather than that of the shoot. In the
germination of many large seeded species, considerable root development occurs, in
sun or shade, prior to emergence of the shoot. Moreover, certain species, e.g. Lotus
corniculatusand Quercusrubra,appear ill-adapted to use their stored materials in height
growth. Nevertheless, several species, e.g. Arrhenatherumelatius, Plantago lanceolata,
and Castanea mollissima are equipped morphogenetically to respond to the diversion
of seed reserve to the shoot.
The susceptibilities that lead to failure in shallow strata of shade may be an indirect
result of adaptations necessary for survival in other environments included within the
recent habitat range (Grime 1965b). Thus failure to escape shade in Arenaria serpylli-
folia, Betula lenta and Betula populifolia was correlated with low seed weight. These
species are colonists of bare soil in abandoned fields and disturbed grassland. For
effective colonization of a habitat which is both transient and small, many seeds must
be scattered. The small seeds which are a liability in shaded environments are, therefore,
a necessity for efficient wind-dispersal onto the rare patches of bare soil.
Several species, including one with comparatively large seeds (Lotus corniculatus),fail
to escape low shade because the seedling cannot form a stable erect growth form. (In
L. corniculatus the internodes expand considerably but apparently do not contain
adequate mechanical tissue to support the etiolated shoot. In turf the surrounding
vegetation may provide some support.) The grassland species in this group (Hieracium
pilosella and Lotus corniculatus)frequently occur in grassland on limestone, chalk and
fixed dunes. Their compact or procumbent growth forms are characteristic of the
vegetation of droughted habitats (Salisbury 1952) and appear to be adapted to conserve
water in that they are growth forms which place the whole shoot in a stratum of relatively
low air turbulence and high relative humidity and, in some species, bring the stomates
close to the soil surface. Furthermore, mutual shading within the rosette causes the
energy absorbed per unit leaf area to be low, reducingthe amount oftranspirationnecessary
to dissipate absorbed radiation. Thus we may postulate that the inability of these plants
to penetrate shade has arisen through adaptation to drought, a factor which plays a
selective role in the present habitats of the species.
(b) Shade tolerance

Seedlings of two grassland species, Deschampsiaflexuosa and Betonica officinalis,were
persistent and showed comparatively low reduction in yield when confined to shade.
This is consistent with their known occurrence in open woods (Clapham, Tutin &
Warburg 1952).
Survival of shaded tree seedlings was greatest in species recognized by foresters to be
tolerant (Table 5). In this investigation tolerance was apparent in trees, e.g. Quercus
rubra and Deschampsiaflexuosa and Betonica officinalis which, as seedlings, have low
growth rates. Clearly this correlation should be tested with more species and later stages
of development. In addition we should see how far slow growth rates limit the success of
tolerant species in unshaded habitats. It is likely that the high growth rates of intolerant
species enable these plants to quickly dominate the vegetation on abandoned land and
waste places. The high growth rates of such species as Rhusglabra and Ailanthusaltissima
are also characteristic of the weeds and ruderal species with which these pioneer trees
compete for survival on exposed soil.

ACKNOWLEDGMENTS
The experimentsat Sheffieldwere carriedout with the support of the Nature Conservancy.
The authors are indebted to Mr David Peckham for his careful assistance in the work at
New Haven and to Dr P. E. Waggoner, Dr G. R. Stephens, Jr., and Dr J. P. C. Kuiper for
their interest in the investigation and their most helpful criticism of the manuscript.
SUMMARY
1. Cylinders may be used to provide shade for comparative study of seedling morpho-
genesis in vertical gradients of light.
2. The facility with which a seedling escapes low shade is related to the amount of
storage material available from the seed and the number and properties of extension sites
in the shoot.
3. Initial height growth in shade is well marked in seedlings of species occurring in
dense grassland (Arrhenatherumelatius, Betonica officinalis,Plantago lanceolata), but it is
negligible in herbaceous species which are restricted to low turf and bare soil (Arenaria
serpyllifolia, Hieraciumpilosella) and in trees which are pioneers of abandoned arable
land (Betula populifolia, Betula lenta, Rhus glabra).
4. Survival of seedlings at low light intensities was greater in species with inherently
slow rates of increase in dry weight. Shade fatalities were more frequent in 'intolerant'
trees, e.g. Ailanthus altissima and Rhus glabra which grow rapidly in full sunlight.
Analysis of the relationship between growth rate and tolerance of shade is complicated
by the vulnerability of intolerant species to fungi.
5. The present investigation suggests that a late phase of forest succession involves the
replacement of species which as seedlings are productive at both high and low light
intensities, by species with slow-growing seedlings capable of enduring long periods in
shade.
REFERENCES
Blackman,G. E. & Black, J. N. (1959). Physiologicaland ecological studies in the analysis of plant
environment.XI. A furtherassessmentof the influenceof shadingon the growthof differentspecies
in the vegetative phase. Ann. Bot. N.S. 23, 51-63.
Blackman,G. E. & Rutter, A. J. (1948). Physiologicaland ecological studies in the analysis of plant
environment.III. The interactionbetweenlight intensityand mineralnutrientsupplyin leaf develop-
ment and in the net assimilation rate of the bluebell (Scilla non-scripta). Ann. Bot. N.S. 12, 1-26.
Blackman,G. E. & Wilson, G. L. (1951a). Physiologicaland ecological studies in the analysis of plant
environment.VI. The constancy for different species of a logarithmicrelationshipbetween net
assimilationrate and light intensityand its ecological significance. Ann. Bot. N.S. 15, 63-94.
Blackman,G. E. & Wilson, G. L. (1951b).Physiologicaland ecological studies in the analysis of plant
environment.VII. An analysisof the differentialeffectsof light intensityon the net assimilationrate,
leaf area ratio, and the relativegrowthrate of differentspecies. Ann.Bot. N.S. 15, 373-408.
Bohning,R. H. & Burnside,C. A. (1956). The effect of light intensityon rate of apparentphotosynthesis
in leaves of sun and shade plants. Am. J. Bot. 43, 557-61.
Bourdeau,P. F. & Laverick,M. L. (1958). Toleranceand photosyntheticadaptabilityto light intensityin
white pine, red pine, hemlock and ailanthusseedlings. Forest Sci. 4, 196-207.
Boysen-Jensen,P. (1929). Studierover SkovtracerresForhold til Lyset. Dansk.Skovforen.Tidsskr.14,
5-31.
Burns,G. P. (1923). Studiesin toleranceof New EnglandForest trees. IV. Minimumlight requirement
referred to a definite standard. Bull. Vt agric. Exp. Stn, 235.
Clapham, A. R., Tutin, T. G. & Warburg, E. F. (1952). Flora of the British Isles. Cambridge.
Grime, J. P. (1965a). Shade tolerance in flowering plants. Nature, Lond. (In press).
Grime,J. P. (1965b).Comparativeexperimentsas a keyto the ecology of floweringplants. Ecology,45 (In
press).

Monsi, M. & Saeki, T. (1953). Ober den lichfaktorin den Pflanzengesell-schaften und seine Bedeutung
fur die Stoffproduktion. Jap. J. Bot. 14, 22-52.
Salisbury,E. J. (1952). Downsand Dunes.London.
Shirley,H. L. (1943). Is tolerancethe capacityto endureshade? J. For. 41, 339-45.
Vaartaja,0. (1952).Forest humusqualityand lightconditionsas factorsinfluencingdamping-off. Phyto-
pathology,42, 501-6.
Vaartaja,0. (1953). Disease caused by weak parasitesassociatedwith shadein seedbeds. Bi-mon.Prog.
Rep. Div. ForestBiol. Ottawa,9, 3.
Vaartaja,0. (1962). The relationshipof fungi to survivalof shaded tree seedlings. Ecology,43, 547-9.
Vaartaja,O. & Cram,H. W. (1956).Damping-offpathogensof conifersand of caraganain Saskatchewan.
Phytopathology,46, 391-7.
Wadsworth,R. M. (1960).The effectof artificialwind on the growthrateof plantsin waterculture. Ann.
Bot. N.S. 24, 200-11.
Waggoner,P. E., Pack, A. B. & Reifsnyder,W. E. (1959). The climate of shade. A tobacco tent and a
forest stand comparedto open fields. Bull. Conn.agric. Exp. Stn, 626, 39.
Wardle, P. (1959). The regenerationof Fraxinusexcelsior in woods with a field layer of Mercurialis
perennis. J. Ecol. 47, 483-97.
WarrenWilson, J. & Wadsworth,R. M. (1958). The effect of wind-speed on assimilation rate-a
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Went, F. W. (1957). The experimentalcontrol of plant growth. Chronicabot. 17, 1-343.
Whitehead,F. H. (1957). Wind as a factor in plant growth. Controlof the Plant Environment(Ed. by
J. P. Hudson), pp. 84-96. London.
(Received 5 November 1964)


Seedling Establishment

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621

SEEDLING ESTABLISHMENT IN VERTICAL GRADIENTS

II. METHODS OF SHADING

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5Ma 1940 15 25 0st 30 e

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FIG. 4. Specimencurvesfor air temperaturesat mid-dayin full sun, August, Connecticut,

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Table 1. Characteristicsof the species in experiment 1

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Days after germination

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Table 2. Yield at 5 weeks of seedlings grown in light gradients 0, 6 and 12 cm

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Table 3. Characteristicsof the species in experiment2

(b) Survival and yield

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Arrowsshow occasionswhen statisticaltests were made of the significanceof differencesof

weight of seedlings after 8 weeks growth in light gradients,

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Table 5. Characteristicsof the species in experiment 3

0 ---------- ______________|-------------------------- ------------ ---

Results: Height of Castaneamollissima, and Quercusrubrawas greater in the unshaded

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1'0 05 0 0'5 10 1-5 2-0 2-5 3-0 35

(b) Survivaland dry weight

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Fatalitiesper container* Maximumheightattained M

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A. Shade avoidance by initial height growth

(a) Source of energy

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(b) Extension sites

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lanceolata (which in closed grassland survives as a rosette of ascending leaves of up to

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(b) Morphologicalresponses increasinglight interceptionin shade

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(c) Metabolic rate

Table 7. Estimated growth rate in full sunlight and mortality in shade

C. Extrapolation of the experimentalresults to thefield

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(b) Shade tolerance

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(Received 5 November 1964)

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