CHAPTER V II

G R O W T H I N R E L A ! 1 0 N

T 0

B I O 1 1 5 F A C T O R S

The biotic factors have since^been considered as

an integral part of the mosaic of factors that control and

influence the growth of a plant species (McDougall 1918).

The biotic factors are important because they w i k as the

directly effecting disjunctive and conductive symbiotic

elements, which influence through association o f species,

pollination, dispersal, competition and parastisim on the

life of a plant species. Clements (1939) stressed » s r the

influence o f biotic factors in three aspects i . e . , •‘action*1

(effect of environment on the organism); "reaction** (effect

of organism on the environment ) and rtco-actionM (mutual

effect between the organisms). In general the biotic factors

have been found effecting the life as also the physical1

surroundings of the plants (Misra 1959), Thus it becomes

rather imperative to study the effect of the various living

organisms (directly or indirectly ) on the plant species*

Owing to the toxic nature o f the white milky

latex present in various plant parts, Buohorbia helioscopia

is rarely giazed by animals though due to their somewhat

diffuse habit the plants are often trampled by man and the

- 153

animals. In addition, becuase of its growth as an unwanted

weed in fields and orchards, it is usually eradicated through

mowing and hand pulling practices. But, as the seeds are the

only means o f propogation, this mechanical eradication

proves effective only during preflowering phase. Eradication

at later stages (flowering or fruiting ) becomes ineffective

as most o f the mature seeds are returned to the soil even

after complete eradication*

Birds have been found to do least damage to the

plants, as they do not seem to have any taste for any of the

plant parts mainly because o f the acrid latex. However, rats

were observed damaging the seed, as under laboratoiy conditions

the seeds kept in open were often eaten away by rats.

The plants are self tBisompatible, as the pollination

in nature is mostly due to flying insects litoe bees, flies,

butterflies and yes& ants. The nectar secreted by the glands

bordering the cyathium forms the chief attraction for these

insects. Two prominent insects namely the green lice (Aphids)

and green moth (Gut worm) have been observed parasitising

the plants. Though the damage cause^by the former is neglegible

the latter insect species has been observed cutting o ff

the entire shoot slightly above the soil at times and mostly

eating away the leaves irrespective o f their size and age

on the plant. The roots of the plant species are often

attacked by nematodes ( Ptylarynchvs %>. ) in damp and

shaded habitats producing innumerable galls on the laterals.

The incidence of the nematode attack is linked with the poor

aeration of the soil. Ealiosome or the o il body used as food

by the ants forms the chief attraction for these insects

to visit the plants. In order to lick away the o il ants

have been observed carrying away the fruits as also the seeds

154 -

from the ground to their respective holes. Late in the

growing season both the seagonal forms o f the plant species

fall pray to rusting disease due to the attack o f Buaomyces

he lioscoplae and Me lams no ra sp. Both the ffcingal plants form

large dark brown to yellow pustules on the leaf surfaces,

more frequently on the lower surface, propcjgating mainly by

te leu to spores. The fungal mycelia form a net woife ramifying

through the intercellular spaces of the leaf mesophylly By

the time the teleutospo res mature, the leaf tissues are

damaged and the leaf growth as also the photo synthetic surface

of the plant reduced. The affected plants 5 riow distinct s i ^ s

of early degeneration under natural conditions of growth. In

the 'summer* form the infection usually starts May onwards

when the plants are at the flowering stage, while in the

•winter* form the attack is most severe during October

to December when the plants are at preflowering stage. Plarits

with and without fUngal infection collected from various

sites were scrutinized for shoot length, average number of

pustules per leaf and the fresh and the dry weight o f the

shoots and the average mean o f 20 readings are set in table 70*

On an average 9 .5 to 10.7 pustules were observed

to occur per leaf surface from the various study sites. As

compared to the uninfected plants the average values for

shoot length, leaf number and the dry weight of the shoot

in diseased plants were observed to be low. The average shoot

length in diseased plants varied from 12.7 to 14 .9 cm in the

•winter* form at various sites as compared to 15.2 to 21 .2 cm

in the healthy plants* The dry weight of the shoot on an

average Varied from 0.412 to 0 .469 gm in the diseased plants

while the values were comparatively higher in the healthy

plants.

- 155 «

Table 70* Effect of rust disease on the growth of

B. helloscooia ( ‘winter’ form).

S. No jstudy site jj&ate o f freight/JNumber o f Jblumoer o f Jury weiglxc fobserva.jj plant jleaves/ Ipustules/tshoot(gm) it ion . Kcm) 8 Plant I plant fi

1. Campus 15 .6 .69 1 2 .7 (1 .2 ? 15(2 .5) 9 .5 (1 .8 ) 0 .412(0 .08)

.d o . 15 .6 .69 15 .2 (2 .1 ) 19(2 .9) 2 0 .586 (0 .12 )

2 . Shalimar 21 .6 .69 14 .6 (2 .7 ) 13(3 .0) 1 0 .2 (1 .5 ) 0 .431 (0 .09 )

-do- 21 .6 .69 15 .8 (3 .2 ) 13(4 .1) - . 0 .592(0.11]

3 . Lal Bagh 12 .7 .69 14 .9 (2 .8 ) 16(3 .5) 1 0 .7 (2 .2 ) 0.469(0.13]

-do. 12 .7 .69 2 1 .2 (3 .9 ) 25 (5 .2 ) » - 0 .875(0 .22)

* ( ) = Standard error

Thus it is evident that the overall growth of the

'winter1 form plants in addition to seed set is hindered

because o f the rust invasion. The ‘ summer1 form plants being

attacked at a very late stage o f growth do not suffer any

damage both ir^erms of growth and seed output.

Under natural conditions o f growth the f"summer*

form plants ;?row well spaced from each other even when forming

pure stands while the 'winter* form plants either occur as

dense stands that are sp*®*d over large areas or in isolated

strips maintaining large or small distances from each other

( P i g 4 2 ) . The 'summer* form plants grow in pure stands during

early phases of growth, which dominate the sites o f their

occurence and prevent the invasion by any other grass or

forb species. The ‘ winter1 form always takes the position

of dominance as in the season of their growth most of the

plant species have already completed their life cycle andUvc

either dried up or died. However, during early spring season

also^the plants of this form dominate the various sites (Fig42).

OOMPSriTIONi

Billings (1957) while reviewing the literature on

competition effects in plants s tat *s that the environmental

and genetic factors are involved during the effect of

competition on survival in times o f stress. He further quotes

n variation in the environmental factors, will determine the

survival of the individuals of the same species of a pure

stand, while the perfectly adapted individuals in mixed

stands will survive in uniform environmental conditions".

Effect of Intra. specific competition on plant growth*

Following experiment was designed to evaluate the

effect of intra-specific competition on the growth performance

of Supho rbia helioscopla.

24 polythelene bags o f 16x12 on size were filled

with garden soil and arranged in two groups o f 12 hags each

for the tw3 seasonal forms. The bags were further arranged

in 4 sets with three jjags per set. Seedlings i*ared from one

year old seeds were transplanted at cotyledonary leaf stage

into the bags according to the following schedule o f densities*

Set No. Number o f plants per bag

1. One

20 Two

3* Bbur

4. Eight plants per bag.

The bags were regularly watered and the plants

were allowed to grow continuously for three months (April to

July ) . Later the plants were extricated from the soil and

scrutinized for Various growth parameters (table 71) fig.

The extent o f growth as measured in terms of shoot

length in the ‘ winter* form was little affected by varying

(*SF = 'summer' form, WF = 'winter' form; ^Significant at

1 percent; ** Significant at 5 percent

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Figure 42* Photographs showing the gorwth per romance ofI . heliogcoad* plants i» different *srt<si&tion? in nature the various slties*

- 158

density while in the 1 summer* form a marked reduction was

observed in the values with increasing plant density. However,

the roots were most affected by competition in both the seasonal

forms as maximum growth of the roots was observed in set 1,

which contained only one plant per bag. The roots in this set

showed an average length of 44,85 cm and 31,5 cm in the ’winter*

and ’ summer' forms respectively, Although the leaf number

per plant did not bear any correlation with the plant density

yet higher leaf dimensions were attained by the leaf on plants

that were grown under lesser densities in both the forms.

In both the forms the growth o f the branches was hindered under

higher densities though branches ' attained variable lengths

in different densities in the 'winter* form. Maximum seeds set

in both the fbrms was observed in set 1, wfeterre the plants

grow in isolation. Plants attained maximum dry matter in both

the forms when grown free of competition effects.

Statistically the values for root length, branch

length and seed output are highly significant at 1 percent

between the various treatments in both the forms, while the

values for inflorescence number per plant and dry weight of the

plants are highly Significant at 5 percent. The growth o f the

two forms maintains a uniform distinction in the different

treatments and various values for the growth parameters of

the plants are highly significant at 5 percent.

Thus from these computations it can easily be

inferred that plants show a difference in their vegetative

and reproductive growth under different densities, their

being an increase in all the growth values under lower

densities of sowing.

n:

The plant species occurs as a dominant in every

type o f association. Thus in order to evaluate the growth

behaviour of the plant species when growing in association

with other plants, 9 different associations were selected

at the University Campus0 The plants dug out from one square

meter area from each of the associations selected for the

purpose were assessed for the growth of the shoot and root

(table 7 2 ) . In these assessments the growth of the shoot

was not seen to be much influenced by the different plant

species that grew in its association as co-dominants or

as mere intruders at the various sites. However, due to

increased competition effects by other snecies as also

their varying densities, the root extension at different

sites varied a good deal.

Eb r survival and quick colonization of the plant

suecies at the various sites during Various seasons^ $he

combination of usually prompt and complete germination with

high rate of natural seeding as also the fast rate of root

and shoot growth, give an edge to the plant species over the

other species, that start growth slightly late.

Reproductive and vegetative growth:

The plant samples collected from different sites

during the various growth seasons were sorted into, leaf,

root and shoot portions. Later these were even oven dried

for determining the weight of the dry matter. The loss in

weight was expressed as percentage moisture in terms of dry

weight of the plant (table ) .

The average values for fresh weight, dry weight,

percentage moisture and the shoot/root ratio of the plant

vary at the different sites (table 73 ) . However, higher fresh

tm lo 9 •"

Interftspeciflc comT

T a b l e ? tr 0 G r o w t h b e h a v i o u r o f 3 . h e H o s c o p i a i n d i f f e r e n ta s s o c i a t i o n s i n n a t u r e .

$ 7 N o jT L i s t o f ~a s s o c i a t e d l e v e r a g e n u m b e r 1 ' l e n g t h 'Ccnfif p l a n t s p e c i e s j o f p l a n t s / X S h o o t j R o o t„L ____ feq* m. . . 6................... _ ! _ _ ______

1 0 P l a n t a g o l a n e e o l a t a 1 5 . 5 + 1 * 2 2 7 . 2 * 2 . 8 1 3 . 7 + 3 . 0TT a f 5 u o u s n a f a n s 2 .0 * * 2 2 . 5 + 5 , 9 3 1 . 5 + 6 . 2TBuifco r b l a rig l i o s c o o i a 1 2 . 0 2 0 . 4 + 3 . 5 2 9 . 9 + 2 . 72 . a a o s e 1 1 a b u r s a - p a s t o r l s 2 1 . 0 2 9 . 5 + 2 . 9 1 9 . 3 + 4 . 2f f u pfao r b l a ^ E e l l o s c o p l a 1 8 . 0 2 1 . 6 + 3 . 7 2 1 . 5 * 4 . 1i—r i m ‘ni n m iw i— — mm■ ■!»i—— »— wtmmm— n— am ^

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2 5 . 4 * 2 . 61 3 . 7 * 1 . °2 9 . 1 + 2 . 3Me d i e a g o s a t j v a 7 . 0 1 2 . 2 * 1 . 8P :SgTOt h e c a s p s " 1 3 . 2 1 3 . 5 * 2 . 03 ."“ "Sg l i o s c o p i a 1 5 . 0 2 5 . 5 + 3 . 0

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3 9 . 2 # - 3 . 5 2 1 , 9 * 2 . 7 1 7 . 5 * 3 . 91 3 . 2 + 9 . 5r u n n e

7 . V e r o n i c a b i l o b ai^r^iioscopir

2 5 . 01 0 . 3 4 5 . 5 + 3 . 72 6 . 5 + 3 . 2 2 4 . 2 * 3 . 52 3 . 7 + 2 . 3

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3 . 0 1 1 . 5 4 . 01 7 . 0 1 9 . 33 1 . 0

2 5 . 5 + 3 . 27 9 . 0 * 3 . 61 4 . 0 * 3 . 23 5 . 1 + 2 . 3 2 6 . 2 + 5 . 5 4 3 . 2 + 5 . 6

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- 161

and dry matter is produced by the ‘ winter* form at all the

sites as compared to the ‘ summer* form*

gm as at site ]& to a maximum of 4,030 gm as at site 3* Out

of this total minimum fresh weight of 0*368 gm per plant,

0 ,218 , 0*130 and 0.020 gm weight are contributed by shoot,

root and the leaf respectively* &nd out of the maximum fresh

weight of 4*030 gm per plant, the shoot, root and the leaves

contributed 3.680 gm, 0 .187 gm and 0 ,163 gm respectively. In

the ’ winter* form out of the minimum fresh weight o f 2 ,195 gm

per plant (site /7^ 1 ,549 , 0 .430 and 0 .225 gm weight are

contributed by shoot, root and leaf fraction of the plant

respectively. Similarly for the maximum fresh weight of

40 .391 gm per plant (site 1 ), the shoot, root and the leaf

fraction conttlbuted 39,084 gm, 0 .950 gm and 0,357 gm to

the total weight o f the plant,

The net biomass calculated in terms of the dry

weight of the plant varies between a minimum of 0 .190 gm

(site 0 to 0 .915 as, the maximum (site 3 ) . Out o f the total

minimum, the shoot, root and the leaf portion contributed

0.062 gm, 0 .040 gm and 0 .007 gm respectively, while for the

total maximum weight, 0 .834 , 0 .049 and 0.032 gm are

contributed by the shoot, root and the leaf portions of the

plant in 'summer* form. In the * winter* form the totalo • u

biomass in terms varies between 0.-494 gm as the minimum

on dry weight basis.

In the 'summer* form maximum shoot/raot ratio

(1 7 .0 ) on dry weight basis is attained at site 3, while in

the 'winter* form the maximum shoot/root ratio of 24 .7 is

In the 'summer* form the average fresh weight

accumulated by the plant varies between a minimum of i>^368'

(at site Jk) to 9 .982 gm as the maximum plant,

- 162 -

attained at site 1,

Total net productivity in relation to growth*

For estimating the total above and under groundp

production for the two seasonal forms fifty 1 m' quadrats

were laid at random at the various preselected sites and an

estimate of the number of plants of the two seasonal forms

in one square meter area was made alongwith the total

number of fruits produced. Monoliths were extricated from

the quadrat sites and after washing away the soil the plants

were sorted into root (under ground) and shoot (above ground)

portions which were dried in an electirc oven to a constant

weight for estimating the dry matter production (table )*

The average density of the 'winter* form varied from

8 .7 plants per quadrat as minimum at site 4 to the maximum

of 29 .35 plants per quadrat at site 11, while in the 'summer*

form the density values varied between a minimum of 9 plants

per quadrat at site 10 to 35 plants per quadrat at site) 3

as the maximum. The values for the reproductive productivity

as measured in terms of total seed out put per quadrat

fluctuated between a minimum of 2205 as minimum at site 3

to 15950 seeds per quadrat as maximum at site 10 in the

•winter* form while the value estimated for the 'summer form

ranged between a minimum of 135 at site 10 to 7484 as

maximum at site 3* The values point to the fact that the

reproductive productivity does not bear any direct correlation

with the plant density at the various sites and is entirely

a function of the plant.

Maximum values for dry natter production of the

under ground parts (4,27lgm per quadrat ) are obtained at

site I in the 'wintetr* form and (2 ,2 7 | gm per quadrat )

are obtained

- 163 -

at site 2 in the 1 summer’ form. The values for above ground

productivity vary between 87*257 gm dry weight per quadrat

and 29.190 gm dry weight per quadrat (site 1 and 4 ) in the

•winter* and ’ summer’ forms respectively as the maximum,

contributed by the shoots. Thus it becomes apparent from

these estimations that the diy matter production bears a direct

correlation with the extent of the vegetative growth of the

plant as si ss ted by the density per unit area*

For the total under ground productivity per

quadrat (a ll the plant species growing therein) 29.457 gm

dry weight at site 10 (0.600 gm ) are contributed by the

•summer’ form plants o f B. heliosoopla. Out of 62.472 gm

dry weight as the maximum contributed by the shoots of a n

the plants per quadrat at site 4, 29.190 gm are contributed

by the shoots of the 'summer’ form*

Thus the average density values workout to be

higher for the 'summer' form while the total productivity

per unit a r e a (reproductive as also the vegetative ) is

considerably higher in the 'winter’ form. This accounts

for the higher survival value and better vegetative growth

of the 'winter' form plants under natural as well as

cultural growth as compared to the ’ summer' form that flower

and fruit immediately after the climatic conditions are

favourable in ^pril to May, due to which the vegetative life

span in this olant form is cut very short.

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Table 74 . Data fo r tha Importance value in d ic ie 1Ks o f the d i f f e r e n t a s s o c ia te s

o f a . h e lio scooja (se a so n a l forms) at d i f fe r e n t study s i t e s .

S.No0 Name o f the____i,plant .spasiqg. F i — r ~ fi

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A n agaH is a rv e n s is

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Artem asia vulgare

A plum

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6 .9

0 .5

4 .5

3 .2

I tA -* -

0.8

2 .3

0 .5

4 .0

1 . 7

10.2 0.8

3 .2

6.2

1 . 4

3 .9

0 .9

4 .2

3 . 1

1.2

5 .2

m

2.2

20.0

3 .7

1 . 9

1 5 .7

1.2

7 . 1

wm

0 . 3

1 2 .4

6 . 8

21.2

1.2

2 .9

4 . 9

Bothreochjtoa 7 . 9 - mm 2 . 8 1 1 . 7 4 . 9 - 5 . 7 mm 2 . 6

Brassica compestris 3 . 8 4 . 1 1 . 5 1 . 7 1 . 2 2 . 5 0 . 8 0 . 7 2 . 8 2 . 8

Broraus scoparius 1 . 5 5 4 . 6 9 . 3 Ml 2 . 4 - 4 . 2 6 . 8 - 6 . 9

Carduous crisp u s 1 . 3 1 . 7 4 . 5 5 . 1 - 2 . 9 3 . 7 2 . 9 - 4 . 7

Carthamus t in e t o m s 2 . 5 1 . 7 - - 0 . 4 wm- 0 . 3 0 . 8 - -

C ap salla b u r sa .p a s to ris 5 . 1 2 . 1 1 . 0 0 . 8 - 2 . 2 3 . 7 4 . 5 1 . 2 1 . 7 2 . 6

Cichorium intybus 4 . 2 1 . 8 - mm 1 . 1 2 . 1 - 1 . 3 - 1 . 7

Convolvulous arvensis 2 . 9 - 1 . 5 mm 2 . 9 0 . 4 - - wm 1 . 0

Ceratocephalus fa lcatus 0 . 2 - mm 0 . 2 - - - - -

Cy,t>3don dactylon 1 5 . 2 1 2 . 9 1 2 . 3 2 0 . 4 5 . 1 1 3 . 4 wm 1 2 . 5 7 O1 O ^ 1 4 . 2

Dacus carotus 1 . 2 1 . 5 0 . 2 0 . 4 0 . 6 - ■■ 1 . 0 - -

19 . Di c ant hium a nnu la turn 15.9 12.2 10.0 6.2 2.7 4.7 5 .1 8.0 2.8 4.520. Erogeron fa lc a tu a 1 .3 - - 2.6 1 . 2 • 0.3 0.42a. E*yangium 0.2 1 . 3 0 .5 » 2 .1 0.8 0.6 mm

22. Euphorbia h e lio scopia(SF) 16 .2 1 3 . 7 2 1 .9 36 .5 2 3 .2 3 1 . 1 19 .4 12 .7 2 5 .2 35 .72 3 . Euphorbia h e lio scopia fw F } 2 1 . 1 1 5 .2 1 3 .8 14 .7 21.2 12.2 15 .6 10*0 3.5.8 21.824. rb£a enjodi 6 . 1 1 . 2 - - 3 .5 1 . 4 1.0 2 .925 . Euphorbia peplus 5 .2 - 2 .1 1 . 8 0 .5 Mi

26. Pumaria p a r v i f lo r a 0 .4 - 0.6 .. 1 . 0 1.2 2 .427 . Heramiun a co n it ifo liu m 0.3 0 .4 0.6 1.2 _ 0 .523 . Oalium aparine 0.1 0.2 wm - 0 .3 Ml 0 . 1 0 .529 . Gnophalium pultoinatum 1 . 2 - vu cs «. 1.2 0.2 wm

30. Hypericum perforatum 1.8 - wm 2.2 Mi 0.2 p. mm M

3 1 . I r t s- - 5 .2 - wm M. wm Ml

32 . K o elp in ia 4 . 1 - - - 1 . 8 0 .9 0.233 . lo tu s c a ra ic u la tu s 4 .2 - 0.9 10.2 5 . 1 1 . 2 2 .7 7 .9 5 .4 4 .234 . Lycopsis a rv e n s is - mm 1 . 8 - 2 .7 - .. wm 0 .335. Marrutoium toulgare 1 . 2 wm m 2 .1 _ 0.8 wm m 0 .436. Medicago d en ticu la ts 1 3 . 1 1 1 . 5 20.2 1 5 .9 7 . 4 3 .2 5 .7 6 .5 1 1 . 4 8.237. Medicaao fa lc a t a 4 .2 5 . 1 - 3 .7 2 .9 1 . 5 mm 5 .2 4 .738. M lig a g o s a t jv a 1 1 . 1 6 .5 1 2 .2 2 .9 3 .2 5 .9 1 0 . 1 5 .0 4 .3 3 .639. O x a iis a c e to s e l la 1 . 4 2 . 1 ~ _ 3 .8 2 .240. JSimSier dutjjyyg 0 .2 0 .3 - _ •» 0.2 o . l Ml

4 1 .

4 2 .

Poa bulbosa 1 . 2 2 . 1 1 3 .2 1 1 . 1 2.0 0 . 3 3 .5 4 .7 2 .5 4 .2

42.

43.

44.

45.

46.

47.

43.

49.

50.

5 1 .

5 3 .

54 .

5 5 .

56 .

5 7 .

58.

59,

60.

6 1 .

32. 6 3 .

Fo lygonum p ie toe gum

Po t t e n t i l a , reptans

Pheleum pratense

Plant a go lanceo la ta

Plant ago ma.jao r

Ranunculii s m c jc a tu s

Rubus 6^ *

S alv ia mo rcroftjana

Set aria g lauca

Sisymbrium as x i llare

Stella rja media

So rghum toulgare

So lanum nig ram

Taraxcum o f f c in a l e

Trigone 11a

T u lipa st e l l a t a

Verba scum thaspus

V lc ia s a t jv a

Veron ica ag re st i s

'Veronica an a g a l j g

Vu lp ia myuros

2.8 •2 . 1

3 . 2 - 1 . 1

5 . 1 1 . 2 1 . 7

1 7 . 1 1 3 . 2 1 5 . 7

CM.10 2 . 3 4 . 9

1 . 2 - 2 . 2

5 . 3 6 . 6 -

5 . 1 2 . 5 1 . 2

2 . 2 4 . 5 5 . 7

oO

- 0 . 8

0 . 3 1 . 2 -

9 . 7 4 . 8 3 . 2

2 . 0 - -

3 . 1 0 . 3 2 . 2

- - 0 . 5

2 . 5 1 . 7 -

- - 4 . 9

o . 4 - 1 0 . 1

0 . 4 - 0 . 1

3 . 2 2 . 3 0 . 2

- 1 . 0 -

I—1 •

3

3 . 2 1 5 .

0 . 8

1.0

2 4 .2

1 7 .2 2 .7

wm

1 . 5

2 .1

0.2

2.8

2 .9

4 .2

1 5 .2

2 .4

1.8

2 .7

4 .2

Mi

4 . 1

0.2

1 . 5

5 .7

1.2

1 . 5

■3.7

4 . 1

2.2

3 .2

10 .7

1 1 . 5

0.2

2 .9

9 .7

1 1 . 9

1 2 . 3

1 5 .2

2 . 2

0 .4

6 .8

0 .8

2.8

3 .0

7 .6

1.1

5 .4

1 . 5

4 .2

2 .5 0 .7

1 1 . 5 7 . 5 2 .8

1.1

2 .5 3 . 3

wm ** ^

1 2 . 3 10 .4 7 ,6

3 .4

1 4 . 3

2 .5

4.6

3 .9

0.6

4 .8

0 .4

2.6

0.9

0 .9

6.2

1.2

11.5