The Relationship Between the Internal Structure and
Photosynthetic Behavior of Apple Leaves

By
William Francis Pickett

A THESIS
Presented to the Graduate School of Michigan State College of Agriculture
and Applied Science in Partial Fulfillment of Requirements
for the Degree of Doctor of Philosophy

Department of Horticulture
East Lansing, Michigan
1935
z '"

ProQuest Number: 10008407

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THE RELATIONSHIP BETWEEN THE INTERNAL STRUCTURE
AND PHOTOSYNTHETIC BEHAVIOR OF APPLE LEAVES

Wm. F* Pickett

Introduction
Several investigators Have made studies of the photosynthet ie
behavior of the foliage leaves of apple (Malus sylvestris) and a few
studies Have been made of the internal structure of leaves of several
species of plants, but the literature does not record any at tenets to
determine if the extent of the intercellular spaces in the spongy
mesophyll of leaves is one of the factors which influence the rate of
photosynthesis•
The purpose of the study Herein reported is to determine
if varieties of apples differ in the structure of the sporgy leaf
mesophyll, if so, the extent of such differences, and if these
possible differences in anatomy have a relationship with variations
in photo synthetic behavior of the leaves*

1

The problem baa three distinct phases of investigation:
(1) the anatomy of apple leaves, (2) stomatal characteristics
and behavior, and (3) photosynthetic behavior*

After the data

on these phases were assembled the influence of the mesophyll
characteristics on photosynthetic activity was studied*

2

PART I.

ANATOMICAL STUDY OP APPLE LEAVES

Historical
Pick (1882) and Stahl (1883) concluded that sun leaves are
thicker than shade leaves, but that the air spaces of the shade leaves
are larger*
de Lamarliere (1892) observed that for equal surfaces, sun
leaves have greater intensity of respiration, assimilation, and
transpiration than those grown in the shade, the well known structural
differences thus having a corresponding physiological significance*
Bonnier (1894) compared the same species of plants at various
elevations in

the Alps and Pyrenees mountains and found that the Alpine

leaves had a better developed palisade tissue, due to larger cells
or an increase in the number of rows, and that there were more stomata
per unit of leaf surface than on 1he leaves of plants grown in the
lowland*

3

Eberhardt (1900) reported tbat humid air favored the production
of larger leaves, and greater amounts of chlorophyll and root development*
Dry air caused the cuticle to became thicker, more stomata to be formed,
and the palisade tissue to be thicker.
Hesselman (1904) studied the influence of light on the leaf
structure of forest trees.

Those leaves grown in the stronger light

had more palisade cells than leaves in the poorer light.

Shade leaves

produced more starch than sun leaves of the same species when the light
is equal.
Bergen (1904) found that sun leaves transpired more than shade
leaves because the greater thickness of the former affords a larger
evaporating surface.
Clements (1904) came to the conclusion that decreased light and
increased water cause an increase in leaf surface and a decrease in
thickness; decreased light causes a somewhat looser arraugment of the
palisade; increased light and decreased water both cause a reduction
in leaf surface and an increase in thickness; strong light causes a
closer arrangement of the palisade; and, finally, that no laws can be
laid down as to the exact amount of change taking place in the histology
of the leaf in response to a definite difference in the physical faetors.

4

Haller and Magness (1925) reported that in order to obtain
apples of good size and quality, from 30 to 40 medium sized leaves
were necessary per apple for Grimes and Ben Davis but for Delicious
even more leaves were required.
Pfeiffer (1928) studied the influences of light of various
wave lengths upon plant growth and found that full insolation produced
thicker leaves than those in the limits of the visible spectrum.
Magness (1928) correlated leaf area and number of leaves with
sugar content and size of apples and concluded that with smaller leaf
area per fruit, the leaves seem to function more efficiently.
Vyvyan and Evans (1932) observed that the size of a leaf varies
with the nature of the growth bearing it, and the number of leaves and
position of the leaves on the growth.
Turrell (1934) computed formulae for the computation of the ratio
of the internal exposed surface to the external exposed surface of
mesomorphic, and succulent leaves.

Though this method may be satisfactory

for comparative determinations for leaves which differ as much as
xeromorphie and succulent leaves, it was not found satisfactory for studies
of varieties of apple leaves.

The differences between the structure of

apple leaves of different varieties are not sufficiently great to permit
the use of this method

5

Pickett (1934) reported that 20 leaves per fruit on Delicious
gave larger fruit than the largest of other varieties with 40 or even
50 leaves per fruit.

The York ratio of 50 leaves per fruit produced

apples which were larger than either Winesap or Jonathan, but smaller
than Delicious at this ratio.

The York trees however, had not produced

fruit the preceding year, but the other varieties had borne.
The weight of fruit produced per 100 square centimeters of leaf
area was determined by Fisher (1934) for McIntosh, Delicious, Rome,
and Newtown.

This vfeight was least with McIntosh and greatest with

Rome.
Chandler (1934) reported fruiting does not reduce wood growth,
in proportion to leaf surface, by as much as the dry matter of the
fruit.

One of the possible causes suggested was that the presence of

the fruit accelerated photosynthesis by removal of the products which
otherwise would accumulate in the leaves and inhibit photosynthesis.

Anatomy - procedure
Seven varieties of apples have been ueed in this study.

The

selection was based on the supposed efficiency of the leaves in
promoting tree vigor and fruit production, rather than upon any previous
knowledge of their internal structure.

Livland and Wealthy were chosen

because the fruit of these varieties ripens from early summer to mid­
summer under Kansas conditions, Livland is rather a dwarfish tree, and
Wealthy trees are of standard size.

(Plates 1, 2, 3, and 4).

Jonathan

and Delicious trees are vigorous, and Jonathan is of major importance
in the Missouri river valley district as a commercial variety.

These

varieties are usually harvested between the first and fifteenth of
September.

Delicious produces such vigorous young trees that they are

extremely difficult to prune properly and wind deformity is a common
characteristic of the variety.
varieties used.

Winesap, Gano, and York were the winter

In central and eastern Kansas, Winesap is one of the

most profitable varieties, especially in those orchards where adequate
cross pollination is provided.

Winesap trees are good in vigor on good

sites but the fruit on the older trees is below medium in size and the
leaves are smaller than those of most other varieties.

-

7

-

Of the varieties

8

Plate 1.

A Livland apple tree, representative

of tliose used in this study.

9

plate

2*

A Jonathan

apple tree, representative

of those used in this study*

10

Plata

3.

A Wealthy apple tree, representative of those used in this

stuiy.

u

Plate 4.

A York apple

tree, representative

of those used

in this study

selected, Gano la one of the most profitable for the canmercial grower.
It Is an annual bearer and the fruit and leaves are usually of good
size*

York is distinctly biennial in its fruit bearing habit*

The

tree is above average in vigor*
The trees in the orchard on the grounds of the Kansas Agricultural
Experiment Station at Manhattan were used for this study*
yearly rainfall at Manhattan is 51*49 inches*
site belongs to the Derby silt loam series*

The average

The soil on this
Considerable soil erosion

has taken place and most of horizon A is gone*

The trees are eighteen

to twenty years old and are fair to good in vigor*

The soil management

method for the past fourteen years has been a combination clean
cultivation, cover crop plan*

Frequent scarcity of soil moisture is

probably the greatest hazard in growing an orchard in this locality, and
this factor becomes of major importance for a period varying from a few
days to several weeks during most growing seasons whan drouth conditions
are aggravated by extremely hot weather and high, hot winds*

During

these periods the evaporating power of the air is great.
Leaves for the anatomical studies were selected from the middle
portions of new shoots on the south side of the outer periphery of the
trees*

Pieces about one centimeter square were cut from near the mid­

section of the leaf blades, only one piece being removed from a leaf,
and plunged in a chromo-acetic acid killing solution containing one

-

12

per cent acetic acid#

The leaf portions were left in the killing

solution over night, after which the killing agent was washed from the
pieces with running tap water*
Dehydration of the samples was accomplished by using the
following series of water, ethyl and N-butyl alcohols as reported by
Zirkle (1930).
Water, cc

95

89

82

70

50

30

Ethyl alcohol, cc

5

11

18

30

40

50

N-butyl alcohol, cc

0

0

0

0

10

20

The samples were allowed to stand for one hour in each step
except the last, which contains a total of 70 per cent alcohol, where
the material was allowed to stand over night.
The dehydration was completed by the following steps*

One hour

was allowed for each except the last, when a 24-hour period was used:
Water, cc

15

5

0

0

0

Ethyl alcohol, cc

50

40

25

0

0

N-butyl alcohol, cc

35

55

75

100

100

Paraffin dissolves extremely slowly in N-butyl alcohol and a simple
method of infiltration is to fill a vial two-thirds full of paraffin,
allow it to harden, put the leaf portions upon it, cover them with N-butyl

13

alcohol and place in the oven.

When the paraffin melts the leaf portions

sink and the N-butyl alcohol stays on the surface because it is lighter
than the paraffin*

The paraffin is changed two or three times and with

the leaf samples used in this study, 18 to 24-hours were sufficient
for the infiltration.

N-butyl alcohol was more satisfactory for this

purpose than xylol in that the infiltrated material was less brittle
and much time was saved*
During the imbedding the leaf portions were arranged with the
lower epidermis toward the bottom of the tray*

A rotary microtome was

used to cut the sections 10 microns in thickness.

The imbedded material

was so arranged on the microtome disc that the knife cut through the
lower epidermis first; the palisade tissue and greater thickness of
paraffin above the upper epidermis prevented tearing of the upper epidermis.
Reversing the direction of the cut sometimes resulted in the spongy
mesophyll and lower epidermis being torn.
A weak mixture of powdered egg albumen suspended in water was used
as the fixative.

After the ribbon was floated on this mixture under

an electric light bulb, nearly all of the mixture was removed by blotting
paper, and the sections thoroughly dried under the overhead heat.

The

staining was done by carrying the slides through the following series:

14

xylol 10 minutes
xylol 1 part, alcohol 1 part, 10 minutes
100 per cent alcohol 10 minutes
95 per

centalcohol 10 minutes

75 per

centalcohol 10 minutes

50 per

centalcohol 10 minutes

2 per cent Safranine 0 in 50 per cent alcohol 4 to 6 hours
50 per cent alcohol 2 minutes
75 per cent alcohol 10 minutes
95 per cent alcohol 10 minutes
100 per cent alcohol 10 minutes
xylol 1 part, alcohol 1 part, 10 minutes
xylol 10 minutes
balsam and cover glass
To permit a study of the looseness or compactness of the mesophyll,
the slides were mounted on a microscope so arranged that it served as a
microprojector*

4 spot light was used as the source of light and the images

from the projector were focused on a screen 43 inches from the outer end
of the microscope barrel so that the leaf sections were magnified
approximately 900 diameters*
Jifty tracings of representative samples of the mesophyll of each
variety were made on paper, not more than five tracings being made of
sections from any one leaf*

15

To secure mathematical descriptions of the relative compactness
of the mesophyll, the cross sectional areas of the intercellular spaces,
as traced on paper, were computed with a planimeter.

By use of a

chartometer the total linear perimeter measurement of each tracing was
determined. These data are presented in Table 1*
During the season 1934, for the orchard studies, only four
varieties were used in contrast with seven varieties in 1933.

The

reduction in the number of varieties was feasible because of the great
similarity between certain varieties and the reduced number made it
possible to determine photo synthetic activity more frequently and
thereby secure more representative records of the daily performance
of the leaves.
and Livland.

The varieties used in 1934 were Jonathan, York, Delicious,
Delicious and Livland were selected because during 1933

they probably had the most compact and the most open mesophyll
structure, respectively, of the varieties studied.

Jonathan and York

were retained because the leaves are large and hence better suited to
photo synthetic studies by the modified Sachs* punch method than the
smaller leaves of Winesap foliage.

16

•

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Table 1.

Intercellular measurements of apple leaf mesophyll

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images, 11 inches long, at a magnification

o
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Measurements for each variety were made from 50 projected
of X900,

S

The differences between measurements of the intercellular
spaces in tne mesophyll of some of these varieties are highly significant
statistically.

Between others, however, the differences are not

significant.
The differences between the means of the intercellular area
measurements for 1933 are presented in Table 2 and their perimeter
differences in Table 3.

As shown by Table 2, the differences between

the cross sectional area measurements of. orchard grown Livland and those
of orchard grown Delicious, Jonathan, Gano, and Winesap are significant
to the extent that the differences are at least four times their probable
errors.

The differences between the means of Livland and the other

varieties vary, in terms of their own probable errors, from 7.8 for the
Livland - Delicious comparison to 3.04 for the Livland - York comparison.
(Picisett 1933j.
Considering the 1934 data, (only four varieties were used) there
was no signif icanee between any comparison of orchard grown Delicious,
Jonathan and York, (Table 4).

However, between orchard grown Livland

and orchard leaves of the other three varieties, the differences in
terms of their own probable errors vary from 3.57 for the Livland York comparison to 2*19 for the Livland - Jonathan comparison.

The 1934

orchard data do not show as great differences in the cross sectional
area measurements as those of 1933.

18

When comparisons between greenhouse grown and orchard grown
foliage are made the differences are greater*

The difference between

orchard grown Livland and greenhouse grown Delicious in terms of its
own probable error is 4.85*

The formula used for calculating the

probable error of the difference between two means is:

* V l _ i% =

V

( ^ v 2 + (KW

19

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Table 2.

Differences between means of cross sectional area measurements of intercellular
spaces in apple leaf mesophyll. Square centimeters. 1933* Orchard grown.

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Livland

Table 3.

Differences between means of perimeter measurements of intercellular
apple leaf mesophyll.
Centimeters. 1933. Orchard grown.

spaces

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Photomicrographs of representative portions of cross sectional
and tangential sections of the apple leaves are presented in
Plates 5 to 14 inclusive.

The cross sections were prepared from

the seven varieties used in 1933.

The tangential views are from

the four varieties studied in 1934.

I'he most important feature of

the tangential sections is the distinct looseness of the Livland
mesophyll when contrasted with that of the York, Delicious and
Jonathan sections.

The tangential illustrations show a portion of

the lower epidermis with a few stomata clearly visible, the Iow &t
parts of some of the palisade cells with the spongy tissue con­
stituting most of the field.

-

23

-

24

Plata 5*

Gross section

(X 200) of a greenhouse grown Livland apple leaf.

35

Plate 6.

Gross section

(X 200) of a greenhouse

grown Delicious

apple leaf

26

Plate 7.

Cross section

(X 200} of an orchard grom

Jonathan apple laaf

87

Plate 8.

Cross section

(X 200) of an orchard grown York apple 1

28

Plate 9.

Cross seetion

(X 200) of an orchard grown Delicious

apple leaf:

29

Plate 10.

Gross section

(X 200) of an orchard grown Livland

apple leaf.

30

Plate 11. Section (X 80), cut parallel or slightly diagonal to the surface of an orchard
grown Jonathan apple leaf* Note stomata at lower left and palisade cells on right.

Plate 12. Section (X 80), cut parallel
grown York apple leaf.

or slightly diagonal

to the surface of an orchard

32

Plat© 13. Section (X 80), cut parallel or slightly diagonal to the surface of an orchard
grown Livland apple leaf. Note stomata at lower right and lower parts of palisade at
top. Compare intercellular spaces in the spongy mesophyll with those in Plate 12.

33

Plate 14. Section (X 80), cut parallel
grown Delicious apple leaf.

or slightly diagonal to the surface of an orchard

PART II.

STOMATAL BEHAVIOR

Introduction

Stomata on apple leaves are found only on the lower surfaces,
and are directly involved in the process of photo synthesis since the
carbon dioxide enters the leaves, for the most part, through them.
Cuticular absorption of carbon dioxide is probably not sufficient
to be of importance in photosynthesis In apple leaves.
Historical
Ho attempt here will be made to give an extensive review of
the literature on stomatal behavior or of methods for making these
studies.

A critical review of the latter together with a bibliography

is given by Lloyd (1908).

He concluded that, if a section of leaf

epidermis be removed and plunged at once into absolute alcohol, the
tissue is dehydrated so rapidly that no measurable change in the
dimensions of the stomata takes place.

34

Sachs had long before pointed

this out to de Tries.

The material thus treated may be studied

in detail at a subsequent time.

Lloyd, working with Fourquieria

splendens and Yerbena ciliata, concluded that stomatal regulation
of transpiration does not occur, except that conservation of the
contained water follows on complete closure of the stomata.

Lloyd

believed that light was the most Important of the environmental factors
that cause stomata to open in the morning.
Loftfield (1921) studied the daily march of stomatal movement
at Salt Lake City, Utah; Minneapolis, Minnesota; and Tucson, Arizona,
and classified the apple with the group of plants, typified by alfalfa,
in which the stomata are closed during the night and open from day
break until midday or until nightfall.

He presents data to show that

stomata regulate the water loss from plants.

The rate of transpiration

is closely governed by stomata when they are nearly closed and by the
factors affecting evaporation when they are wide open.

He also gives a

bibliography on stomatal behavior, especially as it is involved in trans'
piration.

The number and size of the stomata on any given area of leaf

are influenced by the conditions under which they were formed.

A leaf

developed in the shade has fewer and larger stomata per unit than one
produced in li^at.
Maskell (1928) presented evidence to show that diurnal rhythm
of assimilation is closely associated with the diurnal behavior of
stomatal opening.

35

Furr and Magness (1930) found that the duration of stomatal
opening was closely allied with soil moisture content and the
evaporating power of the air.

Favorable soil moisture content caused

the stomata to remain open longer than when the amount of moisture
in the soil is low or near the wilting point*

The period of daily

opening was approximately twice as long on the irrigated plots aa
on the dry plots.

These workers found that there was a close

correlation between the daily period of stomatal opening and the
growth of the fruit*
Penfound (1932) working with the castor bean, found that the
number of stomata varied with the amount of light and indirectly with
soil moisture content.

Sawyer (1932) studied the stomatal apparatus

of the cultivated cranberry, Vaccinum macrocarpon, and found no
chloroplasts in the guard cells.

The stomata never open widely and

the apparatus seemed poorly adjusted to changes in light, temperature,
and moisture*

He reported no differences of taxonomic or functional

significance were found in fbur varieties*

36

Procedure

■Three items involving stomatal studies were investigated in thia
project; namely, the number of stomata per unit area of leaf surface
for each variety of apple used, the size of the stomata as judged by
the length of the stomatal slit or opening, and the diurnal behavior
of opening and closing in a few instances*.
In Table 5 are presented the data on the number of stomata pec
square centimeter of leaf surface for the season of 1933 for both
greenhouse grown and orchard grown trees*

Livland, a Russian variety,

has the smallest number of stomata per unit area of leaf surface for
both the greenhouse and orchard trees, but has the longest stomatal
openiug of any variety used*
In measuring the length of the stomata (Table 5) the opening was
measured and not the length of the adjacent guard cells*

37

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Stomatal counts and measurements were made from strips of the
lower epidermis tarn fran the apple leaves and mounted in water.

At

least twenty-five counts and measurements were made for each variety,
representing one strip from each of twenty-five leaves located on the
south side of the outer periphery of the trees*
were used.

Only mature leaves

Later in this report, data will he presented on the

photosynthetic activity of orchard grown leaves compared with that
of greenhouse grown leaves.

It is of interest to note that greenhouse

grown leaves have fewer stomata per unit area than orchard grown leaves
of the same variety*
Stomata opening and closing was studied for the seven apple
varieties used in 1933 and for the four used in 1934,

No attempt was

made to determine the cause or causes of stomatal behavior, since
the objective of this work was to determine if there might be differences
in this regard among the several varieties.
Two methods of studying the stomatal behavior were employed.

In

one method, the microscope was taken to the orchard and the lower epidermis
of the leaf was examined while still attached to the tree*

This required

a strong light to reflect against the leaf from the sub-stage mirror.
There was much difficulty in using the higher power objectives for this
purpose because of the uneven surface of the leaves and the presence of
great amounts of pubescence on some varieties, especially Jonathan*
Liviand has little pubescence on the lower epidermis.

39

The low power

objective was not satisfactory because of insufficient magnification*
This method was found to be too cumbersome to be satisfactory*

Too

much time is required for each observation and no permanent material is
on hand for subsequent confirmation of the data*
The method which was more satisfactory was to tear off strips
of the lower epidermis and plunge them into absolute alcohol*

Pieces

of the lower epidermis of the leaves were taken by making a small cut
under the epidermis, near the midrib, with a safety razor blade, after
which an assistant quickly jerked with a pair of tweezers as much of the
epidermis as would tear.

The section secured by starting near the

midrib and tearing the epidermis laterally toward the leaf margin, was
triangular in outline because the epidermis tore along the smaller veins.
The epideimal strip was plunged immediately into absolute alcohol, the
entire operation requiring only two or three seconds.

Later the strips

were stained in two per cent safraine 0 in absolute alcohol, rinsed in
absolute alcohol, and permanently mounted in euparal.

The preparation

of permanent slides was necessary because considerable time was required
to determine which stomata were open, due to the fact that with most
varieties the stomata were seldom more than partly open.

It is entirely

probable that with any method of observing stomatal behavior there is
sufficient error to make it practically impossible to determine if a
stoma is closed or only nearly so.

The author has had extreme difficulty

in making determinations of this kind.

40

On August 3 and 4, stomatal behavior was studied on a King
David apple tree at Manhattan, Kansas.

This 20-year old tree was

growing in a heavy clay soil and had been in a wilted condition for
three weeks.

During the few days preceding August 3, the foliage was

wilted during the entire 24-hour period, while during the first part
of the 3-week period, the leaves wilted every day but recovered their
turgidity over night.

The condition of the stomata was determined by

removing strips of the lower epidermis as described above.

In addition

the lower epidermis of the leaves was examined with a microscope without
detaching the leaves from the tree.

At every inspection all of the stomata

were judged to be closed.
The same type of experiment was repeated on August 21 and 22,
1934, with a Jonathan tree.

(Plates 15 and 16).

Bare again all the

stomata were judged to be closed at each inspection.
wilted.

The tree was badly

The data for both the King David and Jonathan trees are presented

in Table 6.

The procedures of judging apparent photosynthesis according

to the modified Sachs dry weight method and of determining the total
acid hydrolyzable carbohydrates are described later in this publication.

41

42
Plate 15»

Wilted Jonathan

foliage,

August

21, 1934,

43
Plate 16,

Wilted Jonathan foliage.

August 21, 1934,

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Stomatal behavior
foliage, 1934.

and photosynthetic activity wilted apple

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Although, for the duration of the experiments, there was a gain
in combustible organic matter with the samples represented in Table 6,
it is considered that it was too small to represent any appreciable
photosynthetic activity and probably falls within the limits of the
error of sampling.

Likewise, the variations in carbohydrate content

are not significant.
Stomatal behavior of greenhouse grown apple foliage was studied
on May 22, and June 5, 1933, and May 2, 1934.

Similar observations on

orchard grown leaves were made on June 12, July 6, and July 10, 1933,
and on July 18, 1934.

(Tables 7, 8, and 9).

No experimental work was

attempted to determine which factors governed stomatal opening and
closing.

Neither were the trees subjected to any special treatment to

prolong the period of stomatal opening.

The primary purpose of the

investigations was to determine whether there were any varietal differences
in stomatal behavior.

It is assumed that stomatal opening and closing

largely governs the rate of photosynthesis in leaves, other environmental
factors being favorable for this process.

A few general statements

summarize these studies:
a.
open

longer each morning than those
b.

open

Stomata on apple foliage in the greenhouse

c.

in the orchard.

With all varieties under observation, many

to wide open until noon in the

usually remain

stomata were partly

greenhouse.

In the orchard, the stomata were seldom open after 8 or

9 o'clock in the morning and on some days they were closed during the

45

entire twenty-four hour period*

There is probably a certain amount of

error in this last statement, because on one or two occasions the leaves
were active photosyntheticaXLy to a greater degree than shown in
Table 6, although the stomata were judged to be closed*

The author

is of the opinion that it i a impossible to determine with certainty
by means of microscopic examination of epidermal strips or of the
attached leaves whether stomata are closed sufficiently to be gas tight
or only nearly closed*
d.

Livland has the smallest number of stomata per unit of leaf

area of any of the varieties observed, but the stomata are longer than
any others*
e.

Wealthy has the greatest number of stomata per unit of leaf

f.

I'ewer Livland and Wealthy stomata are open in the morning

area*

than those of the other varieties*

46

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open and those open to a greater extent; both conditions were listed as being ^opexu*

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PART III.

PHD1DSYNTHETIC ACTIVITY OF APPLE LEAVES

Introduction

No entirely satisfactory method for accurately determining the
rate of photosynthesis has been devised.
obvious.

The reasons for this are

Concomitant with the phenonomenon of photosynthesis are the

processes of respiration and translocation and it is difficult, if
not impossible, to measure the first and exclude the effects of the
latter two.

There are several methods for measuring the apparent

rate of photosynthesis; that is, the amount of photo synthetic
material produced in excess of the amount of substances used in
respiration and translocation.

One of these methods considers the

quantitative determination of the gaseous exchange.
two distinct types of study:

This method has

in one the amount of oxygen liberated is

measured and in the other the intake of carbon dioxide is determined.
The second general method is based on the determination of organic
substance formed or the dry matter accumulated.

50

In the present study, apparent photosynthetic measurements
were made in three ways;

(a) by determining the amount of carbon

dioxide absorbed by a known amount of leaf area, (b) the weight of
the dry matter accumulated per unit of leaf area during a definite
period, (c) comparisons of the total acid hydrolyzable carbohydrates
at different periods of the day.

Each of these methods will be

discussed separately because the results are conflicting.
of them measure the same plant activities.

No two

In each of the three

methods the process of respiration complicates the photosynthetic
determinations*

Not all plant cells carry on photosynthesis, but

all cells involved in photosynthesis also respire.

In photosynthesis,

the carbon dioxide is reduced and oxygen is released, but in
respiration exygen is absorbed, plant foods are used up and carbon
dioxide and water are the end products.
an oxidation process.

Respiration is essentially

Thus, in the cells containing chlorophyll, two

different kinds of chemical processes are taking place simultaneously;
one in which oxygen is released, the other in which oxygen is absorbed.
Since carbon dioxide is utilized in the former and released in the latter,
the amount of this gas absorbed from the outside atmosphere will be less
than the total amount used in photosynthesis.

The process of photosyn­

thesis over any considerable length of time proceeds at a greater rate
than respiration.

51

a*

Determination of the amount of carbon dioxide absorbed by
a known amount of leaf area*

Historical
The method of determining the absorption of carbon dioxide
was first used by Kreusler (1885)*

Other investigators, including

Matthaei (1905), Blackman and Matthaei (1905), Brown and Escombe
(1905), Willstatter and Stoll (1918), Spoehr and McGee (1923),
and Heinicke ard Hoffman (1933) have modified and improved the method*
Good bibliographies on this topic axe found in the works of Stiles
(1925), Spoehr (1926), Miller (1931), Gasaner and Goeze (1932), and
Heinicke and Hoffman (1933)*

Stiles (1925) is of the opinion that

the continuous current method is the most reliable of all methods
that have been evolved for measuring photosynthesis*

In the

continuous current method, a stream of air is drawn past the plant
part under test and through an absorbing solution where the remaining
carbon dioxide is removed from the air*
Any leaf or group of leaves enclosed within a cellophane
envelope, through which a stream of air is drawn is not under entirely
natural conditions.

Miller (1931) and others have pointed out that the

various methods of estimating apparent photosynthesis give conflicting
results because different activities associated with this process are
measured#

It is considered that the intake of carbon dioxide is the

first step in photosynthesis and the increase in dry matter is the last.

52

Miller (1921) points out several details which must be very carefully
manipulated if the results of the carbon dioxide absorption method
are to be of value*

Among these are: (1)

the regulation of the air

stream, (2) the constant temperature of this stream, (3) the carbon
dioxide eontant, (4) the regularity of the illumination, and (5) the
construction of forms of apparatus suitable for the collection of the
gas and at the same time placing the plant under favorable conditions
for photosynthesis*

Heinicke and Hoffman (1933) considered each of

these details and stressed the necessity of using great skill and
accuracy in manipulating the apparatus*

As perfected by Heinicke and

Hoffman, (1933), this method, with proper precautions, gives an index
of relative amounts of apparent photosynthesis under the artificial
conditions to which the leaves are subjected.

These investigators

have attempted to create natural environmental conditions for their
studies*
Procedure
The apparatus used in the series of experiments herein reported
is essentially like that described by Heinicke and Hoffman in Science
(1933).

In this procedure, the author used a series of three absorption

toisers for each day’s run*

One tower was used for a Livland leaf, one

for Delicious, and the third for a blank, through which a stream of
normal air was drawn*

53

For each, determination, the difference of carbon dioxide content
of the air drawn through the blank tower and that of the air drawn
through the towers connected with the leaves was ascertained by titration
with N/10 hydrochloric acid*

Phenolphthalein was the indicator used.

The suction pressure was secured from an aspirator*

The principle on

which the determinations are based is that a N/5 or IT/10 solution of
sodium hydroxide will absorb all of the carbon dioxide from a stream of
air passed through it*
The air current i s divided into small bubbles by means of a
Gooch crucible or a glass crucible with fused-in fritted glass disks*
The sodium hydroxide solution is drawn into a glass tube about 3 cm
in diameter and 75 cm long, supported in an upright position, the crucible
being fastened to the lower end of the tower by a piece of thin walled
rubber tubing.

Below the crucible is a 500 cc suction flask so arranged

that the crucible is fastened to glass tubing which in turn is secured
in the large owning of the flask with a rubber stopper*
of this tube touches the bottom of the flask.
reservoir for the sodium hydroxide solution.

The lower end

The flask serves as a
When the suction is turned

on, the liqaid is drawn up through the crucible into the tower.

The

side arm of the flask is attached to the cellophane envelope enclosing
the leaf under test by means of rubber and glass tubing.
To the top of the tower is attached a piece of capillary glass
tubing so arranged with a manometer that when the rate of air flow is
once determined for a given amcunt of suction, this rate can be duplicated
later by adjusting the flow so that the height of liquid in the two

54

arms of the manometer is also duplicated.
was used to calibrate the apparatus*

A precision wet test meter

During the tests of the rate of

carbon dioxide absorption by leaves, the air was drawn through the
absorption towers at a rate of 2.25 liters of air per square centimeter
of leaf area per hour.
Two hundred fifty cubic centimeters of N/10 sodium hydroxide were
used in each tower in these experiments.

At the end of each run the

solution was allowed to flow back into the 500 cc reservoir and approx­
imately 200 cc of distilled water were used to thoroughly rinse the tower,
crucible, and other fittings.

The dilute solution was transferred to

a 500 cc volumetric flask and about 10 cc of a 25 per cent solution of
barium chloride were added in order to precipitate the carbonates.
volume is then brought up to the mark with distilled water.

The

After thorough

shaking, the flasks are not disturbed until the barium carbonate has settled,
after which 50 cc of the clear solution is drawn off through a pipette, a
few drops of phenolpthalein are added, and N/10 hydrochloric acid is added
until the end point is reached.

The duration of the experiment was five

hours on each of the three days.
The apple trees used in these tests were growing in a greenhouse
and were planted in 12 inch clay pots as one year old budded stock in
January, 1934.

The tests were made in August, 1934.

55

At the "beginning and at the conclusion of each experiment,
leaf punches were taken from 50 leaves of each variety and used as
an index of photosynthesis by the dry weight accumulation method.
Also entire leaves were collected at these periods, killed by dry
heat in an electric oven, and later analyzed for total acid
hydrolzyable carbohydrates, expressed as glucose.
for these methods is described later.

The procedure

This plan makes it possible

to compare these three methods of judging apparent photosynthetic
activity of apple foliage under the same environmental conditions.
These data are presented in Table 10.

There is no consistency in

the results of the three methods, and it would be difficult to select
one method as the standard of comparison, by which to judge the
accuracy of the other two.

Heinicke and Hoffman (1933) found that

there was great variation in the rate of photosynthesis shewn by
adjacent leaves on the same shoot, also on different shoots of the
same variety.

It seems justifiable therefore to expect to find different

photosynthetic values for leaves on different shoots or even different
trees of one variety as well as between leaves of different varieties.

56

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70

of greenhouse grown foliage, August 15, 16, and 17, 1934.

(0
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Table 10.

Apparent photo synthetic activity

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The same two leaves were used for all the experiments.

The area

of the Delicious leaf was 39.49 cm2 and that of the Livland leaf, 40.09
cm • At the conclusion of these experiments, the two leaves were removed

from the trees and mounted sections for microscopic study were prepared,
as described before.

(Table 11).

The ratio of the average weight of carbon dioxide assimilated
per square meter of leaf area for the three 5-hour runs to the average
weight of the total acid hydrolyzable carbohydrates accumulated per
square meter of leaf area for the same periods is 8.59 for Delicious
and 8.58 for Livland.

However, the ratio between the increase of the

weight of the dry matter per square meter of leaf area and the
carbohydrate gain for Delicious is 11.09 and for Livland it is 5.62.

The

small gain in dry matter recorded for Livland on August 16, is no doubt
responsible for part of hie discrepancy between these two ratios.

The

total molecular weight of the six molecules of carbon dioxide entering
into the formation of each molecule of glucose is 264, and the molecular
weight of glucose is 372.

The ratio between these molecular weights is

1.409, which does not agree with the ratios of 8.59 and 8.58 above.
The photosynthetic activity of these leaves is greater when judged
by the dry weight accumulation than by the carbon dioxide absorption
method.

All three methods, give higher values for the Livland leaf than

for the Delicious leaf, and it is significant that the intercellular
suaces in the Livland leaves are more extensive than those of Delicious.

58

Table 11* Intercellular measure me nt s of tlie mesophyll
of apple leaves used with carbon dioxide absorption
towers in greenhouse, August 15 to 17, 1934**

Variety

Delicious
Livland

Average cross sectional area
of intercellular spaces, cm

Average perimeter measurements
of intercellular spaces, cm

87*01+7.01

179.78+14.21

107.71+12.19

192.79+16.06

* Measurements for each variety were made from 50 projected images,
11 inches long, at a magnification of X 900*

59

b,

The Dry Weight Method,

The weight of the dry matter accumulated per unit of leaf area
is the second method used in this study of the photo synthetic activity
of apple leaves.

Historical
Sachs (1884) was the first to use the dry weight method of
measuring the rate of the formation of carbohydrates.

He removed one-

half of a leaf blade at the beginning of an experiment and left the
other half attached to the midrib,

Portions of the severed half were

measured for area and the dry weight determined.

The half leaf left

on the plant was removed at the end of the experiment, measured, and
weighed as before, care being taken to avoid large veins in both cases.
The gain in dry weight per unit area of leaf surface was attributed
to the carbohydrates formed by photosynthesis,

Sachs recognized that

translocation and respiration should be measured and discovered that
detached leaves showed a greater gain when illuminated than leaves which
remained attached to the plant,

Ha attributed the smaller gain in

the latter case to the translocation of the products of photosynthesis
to other plant parts.

In order to obtain the corrected photosynthetic

value therefore, he added the loss in dry weight per unit area during
the night to the gain in dry weight per unit area found during the day.

-

60

The dry weight method has been criticised by several investigators.
Brown and Escomb (1905) compared the results of measuring photosynthesis
by the continuous gas stream method and the dry weight method.

Working

with Catalpa bignonioides they recorded values two and three times
greater for the dry weight method than that calculated from the amount
of carbon dioxide absorbed.

They concluded that the great objection

to Sachs* method is that all of the errors become accumulative in the
final result.

Table IE is a partial summary of their study.

61

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Ganong (1908) was not convinced that the dry weight method gave
results much too high*

He designed (1905) a punch which could be used

to remove circular discs, having a diameter of 1 .12Q cm or an area of
1 square centimeter, from leaves.

Since that date, most of the dry

weight records have been made by using the punch, rather than the
half leaf method,

Ganong (1908) concluded that the removal of the discs
i

*

does not interfere seriously with the functioning of the leaves.
Too little is known of the composition of the products of
assimilation to make possible an accurate calculation of the carbon
dioxide absorbed from a given increase in dry weight, (Thoday, 1909).
He recommends that the ash content of the leaf samples be determined
and deducted from the increase in dry weight.

According to this

investigator there are two main errors in the dry weight method.
arej

(l)

These

the shrinkage of leaves in area during the experiment, and

(2) their lack of symmetry in respect to dry weight per unit area.

The

latter objection may be overcome by avoiding large veins and using a
large number of leaves for each experiment.

Thoday concluded that if

the dry weight gains are as great as 2 milligrams per square decimeter
per hour, the method is capable of yielding useful results for comparative
studies of assimilation.

In 1910 he found that Helianthus leaves increased

in dry weight by 17 mg 100 cm2 hr., at a time when catalpa gained only
5 mg. 100 can2 hr., a fact which is correlated with the absence of stomata
from the upper surface of catalpa leaves.

65

Spoehr (1926) sums up the merits of the dry weight method by
stating that it offers the simplest means of estimating photosynthesis
and that its very simplicity makes it highly desirable, but in its
present form it is not reliable.

He emphasizes the necessity of

correcting the values for respiration and translocation.
Miller (1931) has used the dry Height method for making
photosynthetic comparisons between, corn, milo, and the sorghums under
field conditions in the Great Plains.
The dry weight method was used more extensively than any other
in these photosynthetic studies because it is the best suited to
orchard conditions.

Whatever may be said against this method, it remains

that the results are comparable, and in the final analysis no method
is accepted as absolutely accurate.

Every possible precaution to avoid

error was adopted in planning these experiments.
precautions are:

Chief among these

(1 ) many of the samples were corrected for variations

in ash content, (2) large numbers, fifty or one hundred leaves were
selected for sampling each variety for each experiment, (3) careful
measurements showed that the leaves did not shrink in area during the
daylight hours, (4 ) dust, spray residues and similar material were washed
from the leaves twenty-four hours before the beginning of each experiment,
(5) in the removal of each disc with a Ganong punch, care was exercised
to avoid the cutting of large veins, (6) comparable leaves were always
selected, only mature leaves on new shoot growth being used, (7) no leaf
showing injury caused by any fungus, insect or other pest was sampled,

64

(8 ) all leaves from which, discs were cut were located on the south
and southwest exposure of the outer periphery of the tree and were
located at heights' ranging from four to six feet from the surface of
the ground, (9) none of these leaves was at any time between the hours
of 5 a.m. and 2 p.m. shaded by the foliage of other trees, (10) all the
trees were growing under practically the same method of soil management.
Plate 17 illustrates the relative sizes of representative leaves
of the varieties under test*

Most orchard grown apple leaves are not

sufficiently large.to permit the removal of more than six discs each
having an area of one square centimeter without severing large veins
or the midrib.

Accordingly many of these experiments were so planned

that only four or six discs were removed from each leaf.

Several

preliminary tests revealed that apple leaves in the orchard ceased to
gain in weight about midafternoon, that is, the processes of respira­
tion and translocation then proceed at a faster rate than does apparent
photosynthesis.

The leaves in Plate 17 were pressed between two pieces

of glass to aid the photographer to secure a sharp focus.

The pressing

so flattened the leaves that the holes do not appear to be circular,
but in natural position the holes were round.

65

66

Plate

17. Representative leaves (X 5/8) with punches removed:
(1) Jonathan,
(2) Winesap, (5) York, (4) Gano, (5) Delicious, (6) Wealthy, and (7) Livland,

In January, 1933, two lots of one year old apple trees were
planted in greenhouses in 12 inch clay pots.

One lot of trees were

plunged in a ground bench in a house designated as Plot A, where the
temperatures were kept as high as the heating facilities permitted.
The day temperatures usually ranged from 90°F to 110°E and the night
temperatures were seldom below 70°E.

Effort was made to keep the

relative humidity high in this house, by keeping one aisle flooded
with water and at times, a small stream of water was directed on the
steam heating pipes.

The other lot of trees, Plot B, were plunged

in a ground bench in a house largely devoted to the culture of lettuce.
The day temperatures were kept at about 70°F or slightly lower, and
the night temperatures were in the upper forties.

The object of growing

the trees under these two sets of environmental conditions was to
attempt to produce within a variety leaves with variation in the extent
of intercellular spaces in the spongy mesophyll.

The two greenhouses

were adjoining and on the days the leaf punches were collected the
door between the houses was open, and the temperatures and relative
humidity were kept the same in the two r ooms.

In Table 13 are presented

some data of anatomical and photosynthetic studies on leaves of these
two groups of trees*

67

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Table 14*

Temperature and relative humidity records in greenhouse*

Date - 1933

April
5

April
6

April
26

April
27

Temp eratu??e (G .)
Relative liumidity
per cent.

15

8

10

16

66

58

56

68

T emp er ature (0 .)
Relative humidity
per cent*

27

22

20

34

45

45

42

39

15

15

11

21

61

63

58

50

T ime of day
7•oO a.m.

12: 00 Noon

5:00 p.m.

Temperature (Ci.)
Relative humidity
per cent.

69

Many factors are blended "together in governing the rate of
photosynthesis and in determining the quantity of photosynthate produced*
It is extremely difficult to select any one of these factors and attempt
to demonstrate the definite role it occupies in a process as complicated
at this*

As one variable is changed, its effect on the rate of photo­

synthesis is masked by the overlapping influences of the other variables*
Nevertheless, the data in Table 13 indicate certain trends of behavior between
the two variables of leaf structure and photosynthetic activity.
(a)

These are:

the leaves of Livland, Jonathan, and York grown on trees in Plot A

have a greater extent of intercellular spaces in the spongy mesophyll than
leaves of the corresponding varieties grown in Plot B, where relatively
low temperatures prevailed when the leaves were growing, while the reverse
was true with Gano; (b) within each variety, the leaves possessing the more
extensive intercellular spaces produced the greater gain in dry weight per
square meter of leaf area between 8:30 a.m* and 5:30 p.m.; (c) the inter­
cellular spaces in the Livland mesophyll from Plot A are larger than those
of other varieties in the same plot.

The same is true comparing Livland

with the other varieties in Plot B, (d) Livland leaves accumulated a greater
average weight of dry matter per square meter of leaf surface than the other
varieties within each plot.

I

70

These records do not isolate leaf structure from the other factors
which regulate photosynthesis, but it is likely that variations in
intercellular spaces partially influence this activity or accompany it
under the dominance of other more powerful factors.
Air temperature and relative humidity data in the greenhouse
for April 5, 6, 26, and 27, 1953, are presented in Table 14.

In

Table 15 the percentage of water In the apple leaves used in photo­
synthetic studies on the above dates is given.

71

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Scar city of available soil moisture accompanied by high,
■temperatures constitutes one of the hazards of crop production in the
Great Plains*

The season 1933 in Kansas was characterized by high

temperatures and below normal rainfall*

Soil moisture determinations

were made on 10 days between June 14 and August 23*

Between June 14,

and July 7, the soil moisture content in non-irrigated plots ranged
from 15 per cent to 17 per cent which is not far above the wilting
coefficient*

Although the apple trees used in these studies did not

exhibit wilted foliage, 1400 gallons of water were used to irrigate
each tree*

Daily maximum temperatures exert a pronounced influence

on the amount of dry matter gained per unit of leaf area, according
to Plate 18*

Generally, the higher the daily maximum temperature, the

lower the leaf gain in dry weight.

Soil moisture data are not included

in Plate 18 because it was impossible to collect representative soil
samples in the vicinity of the greatest root concentration, due to the
irrigation given each tree*
Bose (1924) concluded that the rate of photosynthesis is the
expression of the combined effect of the factors of light and temperature*
Photo synthetic activity increased from 7:30 a.m. to 12:00 noon.

After

2:00 p.m. both light and temperature underwent a decline, with
resulting rapid fall of activity.

His findings are in accord with the

relationship between temperature and photosynthetic activity as shown
graphically in Plate 18.

Sunshine prevailed on each day with the

73

PLATt

18

RELATIONSHIP BETWEEN MAM/AU/A AIR TEMPERATURE
ANDTOTAL DPT WEIGHT GAIN PER 5Q.UARE /ALTER
LEAf AREA E0 R SEVEN APPLE VARIETIES
1933

74

exception of August 25, which was cloudy.
intensity were made*

No records of light

Considering only the maximum air temperatures

on the eleven days when the dry weight determinations were made these
records show the depressing effect of extremely high temperatures on
photosynthetic activity.
An experiment was designed in 1934 to compare the efficiency
of greenhouse grown and orchard grown leaves of Livland and delicious
apple leaves under orchard conditions (Pickett, 1934).

It has been

shown in Table 1 that there are marked differences in the extent of
the intercellular spaces in the spongy mesophyll of greenhouse grown
Delicious and orchard grown Delicious leaves.
exists with Livland leaves.

A similar condition

In January, 1934, several one year old

whips of the two varieties were planted in 12 inch clay pots and
placed in a greenhouse.

In July, 1934, these trees were taken to the

orchard, and the pots were plunged in the orchard soil for a few days,
to bring the soil moisture in the pots to approximately the same
point as that in the soil outside the pots.

The photo synthetic

determinations of the greenhouse grown trees could then be made under
the same environmental conditions as the orchard trees.
This experiment was started on the last day of July, 1934, a
month which broke all records at Manhattan for the past 75 years for
high temperatures of any month.

On twenty-six days the maximum

temperature was 100 degrees F or higher and during eleven consecutive
days the maximum reading was 111 degrees or higher.

75

The rainfall for

"the month was 0#86 inch, and at the time of this experiment the
average moisture eontent of the top first, second, and third foot
samples of soil was 11*6 per cent, dry'weight basis#

Many trees in

the orchard were wilted but the ones used in this test of July 31
to August 1, were apparently turgid#
The rate of apparent photosynthesis was judged by Ganongra
modification of the dry weight method*

Discs one square centimeter,

in area were punched from 50 mature leaves, one disc being removed
from each leaf at 6:00 a#m#, another at 2:30 p*m., a third set at
6; 00 a. m. the second day.
In Table 16, data are presented to show the gains and losses in
dry weight per square meter of leaf area from 6:00 a#m#, July 31,
to 6:00 a. m . , August 1, 1934#

The orchard grown Livland leaves made

a net gain of 5#93 gms M 2 leaf area while the greenhouse grown Livland
leaves gained 3#48 gms M 2 leaf area#

Likewise, the orchard grown

Delicious leaves gain more in dry matter than those grown in the
greenhouse#

These gains were 4#68 grs

area, respectively#

leaf area and 1.88 geos M 2 leaf

The four groups of leaves rank as follows in the

net day gains in dry weight: (l) orchard grown Livland, (2) orchard
grown Delicious, (3) greenhouse grown Livland, and (4) greenhouse grown
Delicious.

The same ranking applies to the measurements of the

intercellular spaces in the mesophyll as shown in Table 1#

Apparently

the differences of the extent of the intercellular spaces are reflected

76

in the photosynthetic behavior of the two varieties*

Several factors

enter into and govern the rate of photosynthesis and the extent of
the surfaces of the exposed cell walls bordering the intercellular
spaces is probably one of them*

7?

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Temperatures;

Table 16*

Variation in dry matter of apple leaves

O

July 31, 23
degrees C relative humidity 42
July 31, 38
degrees G relative humidity 21
Aug. 1, 28
degrees C relative humidity 37
cloudy, practically no direct sunlight at

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Dry weight determinations of orchard grown apple leaves were
recorded on these

dates in 1933:

June 7, June 14, June 17, June El,

June 23, June 29,

July 1, July 6, July 7, July 10, July 25, August

3,

August 9, and August 25.
Photosynthetic activity as measured by determining the amounts
of total acid hydroloyzable carbohydrates expressed as glucose was
determined on July 10, July 25, Augjust 3, August 9, and August 25,
1933.
In 1934, both dry weight and carbohydrate determinations were
made from samples

collected May 2, May 16, May 23, June 5, July 2,

July 3, July 9, July 10, July 18,

July 19, and August 3,August 6,

August 7, August 15, August 16, August 17, August 21, August 22,
August 23, and August 24.

The data for the two seasons are presented

in Tables 16 to 41.
c.

Photosynthetic Activity as Judged by the Saccharification Method.
In this procedure, the entire carbohydrate content is

hydroloyzed to hexoses and the products thus formed are estimated as
glucose by the reduction of Fehling*s solution.
these studies follows:

The method used in

Weight 1.5 gm of dry finely pulverized leaf

powder, add 150 cc 2|- per cent hydrochloric acid, boil in water bath
under reflux condenser for two hours; cool, nearly neutralize with
sodium hydroxide solution, clear with a saturated solution of neutral

79

lead acetate solution and filter.

Add sufficient solid sodium oxalate

to precipitate all the lead and refilter through a dry paper into
S50 cc volumetric flask.

Test the filtrate for the presence of lead

with a little solid sodium oxalate and filter if precipitate forms.
Bring to volume with distilled water.
Prepare Soxhletfs modification of Fehling’s solution hy mixing
equal volumes of (a) and (b) immediately before using,
(a)

Copper sulfate solution - dissolve 34.639 gms CuSo^.5

HgO in water, dilute to 500 cc and filter through prepared asbestos,
(b)

Alkaline tartrate solution - dissolve 173.0 gms of

Rochelle salts (sodium potassium tartrate) and 50 gms of sodium
hydroxide in water, dilute to 500 cc allow to stand for two days, and
filter through prepared asbestos.
Transfer 25 cc each of the copper sulphate and alkaline tartrate
solutions to a 300 cc Erlenmeyer flask and pipette exactly 50 cc of the
sample containing the hexoses and heat in hot water bath at 80° centigrade
for 30 minutes (Quisumbing and Thomas 1921).

Remove, cool and add

25 cc of a potassium iodide-iodate solution (Shafer and Hartman,
1920-1921) prepared by dissolving 60.0 gms of potassium iodide and
5.4 gms potassium iodate in water, adding a few cc of concentrated
sodium hydroxide and diluting to 1 liter.
16 cc of 5 U sulfuric acid.
is dissolved.

Quickly add approximately

Shake the flask until all the copper oxide

Next add 20 cc of a saturated solution of potassium

oxalate and titrate with a standard N/10 solution of sodium thiosulfate

80

to which has been added a small amount of concentrated sodium
hydroxide.

Record the number of cc of the sodium thiosulfate

solution required to oxidize the iodine and substract this from
the amount used when 50 cc of water is substituted for the sugar
solution.

The addition of a small quantity of soluble starch shortly

before the end point is reached makes the end point more distinct#
The net titration gives the amount of sodium thiosulfate which has been
used to reduce the iodine vhich had been used to oxidize the cuprous
oxide and if multiplied by the factor 6,36 will give the number of
milligrams of metallic copper reduced by the sugar#
The acid is added as directed above to release free iodine
from the iodide-iodate solution#

The potassium oxalate is added to

unite with all the divalent copper.

The iodine oxidizes the cuprous

copper to cupric oopper and the sodium thiosulfate reduces the iodine.
The tables of Quisumbing and Thomas (1921) were referred to for cal­
culating glucose frcan the copper value#
It is important to standardize the sodium thiosulfate solution
as follows;

Prepare a N/10 solution of potassium biiodate.

Pipette

exactly 50 cc of this solution into a 250 cc Erlenmeyer flask and add to
this a solution of 3 gms of potassium iodide dissolved in 25 cc of water.
Then add 10 cc of 5 II sulfuric acid and titrate with the 11/10 sodium
thiosulfate solution.

If the number of cc of biiodate used divided by

the number of cc of thiosulfate used is more or less than one, the
quotient secured is a factor by which all determinations must be
multiplied#

81

The leaves used in these analyses were killed in dry heat
in an electric oven.

They were placed in convenient trays made of

hardware cloth and for the first hour the oven temperature was 70°
C.

During this hour the door was left open slightly to permit the

ready escape of water vapor.

The killing was completed at 98° C

after which the leaves were ground with a mortar and pestle until
the powder would pass through a 40-mesh sieve*
a powder which was a natural green.

This method produced

If samples of leaves were put

in the oven at 98° without first going through the 70° step, generally
the powder was somewhat darkened.
methods, were made.

Ho comparative analyses of the two

It was assumed that the bright green powder was

the better.
None of. the trees used in 1933 had apples on them..

Trees not

in bearing were selected because the presence of fruit probably
influence the photosynthetic activity of nearby leaves.

In 1934,

the photosynthetic behavior of leaves on branches bearing apples was
deterniined.

For comparison, leaves on branches from which the fruit

had been removed were used.

The ratios given in Tables 35, 36, 40,

and 41 indicate the number of leaves per fruit on the branches from
which the leaves were selected for study.
In Tables 25 to 41 inclusive the photosynthetic activity of
apple leaves is presented, based on both the dry weight and saccharification methods.

82

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Variation in dry matter and water content of apple leaves

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Variation

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of apple

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to
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o>

Livland

Table

30.

Variation

in dry matter and water content

of apple leavea

-rH

-p
pi
<
u
8 S

CO CM

3•3 .
C
M
O

+

+ I

+

I

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ss

o

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tO H tO
H. W. .O

rt Pi

o

to ^

o

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IN

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to o

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to
to sit

...

CO
to to LO

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to to to

o

CM CM CM
H tO CT*

to

to to to

CM CO

0> r-l

CO o

3•$•
C
O
C
M

lO C
M

+ I

+ I

+

.

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I
.

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i

o» c>- o
to CO

OC
t
- tO
C
DO ^

tt)O H
H H ts

CO C- CO

0 0*0

C
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D

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to to to

to to to

o

CO O
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•

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to to to

+ 5.38
- 4.98

tO Q
lO
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62.46
61.82
62.46

<D

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72.30
77.68
72.70

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355

Livland

Table 21.

Variation in dry matter and water content of apple leaves

tu

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09 i—I
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H
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CM O SO
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50 tO to

to o

to

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to to to

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to to to

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66.20

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61.59
63.02

13
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88

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Livland

Table 22.

Var iation

in dry matter and water content of apple leaves

o

GO to
O GO
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4- 4-

+ 3.94
- 1.46

•
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to o>

Table

24.

Variation

in dry matter

and water content of apple leaves

H 4 CM CO

H* to
rH O

d>

th

* • • • • • • • «

« • • • • • •
a a a a a aa a a a aa a a aa a

9
,a

R

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a a a a a a a a

• * • • » • * •
• • • * • «
(0 ft <0 ft <0 ft (0 pi <0 ft id ft aJ ft <d ft (d ft <0 ft (0 ft Cd ft <0 ft <0 ft
o OOOo o o o o
O o
o o o
o o o O o o
o
O o O
o o o o o o o o o
Oo o o 8 o o o O o o 8 O o 8 o o o
in m in in m o V
CN
n
m
i
)
>
D
in
I
N
in
in
m
in
I
N
i
n
INin o- in o
• • • •

O

• • • • •

B
S1
m

Mft
£

s

Ha

90

o

8
(0
c!>

■rH

rH

®

&
r

■d
£

CO
I
—I
rH

Table 25. Analysis of apple leaves*
July 10 to 11, 1933* Variation
in dry matter, water content, and acid hydrolyzable carbohydrates
estimated as glucose*

Variety

Jonathan

Winesap

York.

Grano

Delicious

Wealthy

Livland

Grams per square meter leaf
Dry
Gain or Carbohy­
matter •
loss of drates.
total
dry
matter.

Time

5:00
10:00
1:00
4:00
7:00
5:00
5:00
10:00
1:00
4:00
7:00
5:00
5:00
10:00
1:00
4:00
7:00
5:00
5:00
10:00
1:00
4:00
7:00
5:00
5:00
10:00
1:00
4:00
7:00
5:00
5:00
10:00
1:00
4:00
7:00
5:00
5:00
10:00
1:00
4:00
7:00
5:00

a.
a*
p*
p*
p.
a*
a*
a*
p.
p.
p.
a*
a.
a*
p.
p.
p.
a*
a*
a.
p.
p.
p.
a.
a.
a*
p.
p.
p.
a.
a.
a.
p*
p.
p.
a*
a*
a.
p.
p.
p.
a*

m.
nu
m.
m.
m.
m*
m.
nu
m*
nu
m*
m.
m.
m.
nu
m.
m.
m.
m*
m.
m*
m*
m.
m*
nu
m*
m.
nu
nu
m.
nu
m*
m*
m*
nu
nu
nu
nu
m.
nu
nu
nu

65.88
71.20

+ 5.32

70.68
64*78
80.66

- 0.52
- 5.90

89.34

+ 8.68

86.58
83.48
63.72

- 2.76
- 3.10

69.08

+ 5.36

69.60
68.48
70.60

+ 0.52
- 1.13

74.16

+ 3.56

76.22
73.60
83.98

+ 2.06
- 2.62

89.36

+ 5.38

90.74
87.68
73.06

+ 1.38
- 3.06

80.48

+ 7.42

78.32
82.06
65.88

- 2.16
+ 3.74

71.20

+ 5.32

70*68
64.78

- 0.52
- 5.90

91

9.16
10.94
10.72
12.51
13.61
11*47
13.03
13.62
14.06
14.91
13.75
10.83
10.88
11.03
11.28
14.66
13.42
11.05
11.30
14.52
11.87
14.71
10.35
9.52
9.85
9.78
8.79
9.05
11.61
10.80
10.93
11.58
13.89 '
13.48
12.44
10.62
8.44
8.63
10.69
10.44
10.15
9.49

area.
: Percentage•
Gain or
Water.
loss of
carbohy­
drates*
60.21
+ 1.56

54.05

+ 2.89
- 2.14

55.12
64*08
58.15

+ 1.03

51.54

- 0.31
- 2.92

53.55
63.83
61.23

+ 0.40

55.93

+ 2.14
- 2.37

57.20
66.92
57.14

+ 0.47

53.50

- 1.52
- 0.83

54. 64
61.44
58.41

- 0.06

54.73

+ 2.62
- 0.81

55.90
61.28
59.48

+ 2.96

56.21

- 1.45
- 1.82

58.09
60.78
61.07

+ 2.25

58.64

- 0.54
- 0.66

60.41
63.40

Table 26. Analysis of apple leaves. July 25 to 26, 1933. Variation
in dry matter, water content, and acid hydrolyzable carbohydrates
estimated as glucose.

Var iety

Jonathan

Winesap

York

Gano

Delic ious

Wealthy

Livland

Grams per sauare meter leaf
Dry
Gain or Carbohy­
matter •
loss of drates.
total
dry
matter

Time

6j 00
10i 00
Is 00
4j 00
7i 00
6s 00
6!,00
10s 00
1; 00
4;00
7:00
6:00
6s,00
10s 00
Is 00
41,00
7:00
6s,00
6:00
10:00
1: 00
4:00
7siOO
6s,00
6,.00
10:00
1 00
4 00
7 00
6 00
6 00
10 00
1 00
4 00
7 00
6 00
6 00
10 00
1 00
4 00
7 00
6 00

a.
a.
P*
PP.
a.
a.
a.
PP*
p.
a.
a.
a.
P*
P«
P*
a.
a*
a.
PP*
P*
a.
a.
a.
P*
P*
Pa.
a.
a.
P*
P*
P*
a.
a.
a.
PP*
P*
a.

m.
m.
m.
m.
m.
m.
m.
m.
m.
m.
nu
m.
m.
m.
m.
m.
m*
m.
m.
m.
m.
m.
m.
m.
m.
m.
m.
in.
m.
m.
m.
m.
m.
m.
m.
m.
m.
m.
m.
m.
m.
m.

81.60
86.54

+ 4.94

86.20
89.82
91.78

- 0.34
+ 3.62

95.74

+ 3.96

96 #70'
94.12
71.16

4 0*96
- 2.58

75.44

+ 4.28

75.14
76.00
79.42

- 0.30
+ 0.86

85.20

+ 5.78

84.86
84.48
82.14

- 0.34
- 0.38

90.32

+ 8.18

86.56
87.76
89.04

- 3.76
+ 1.20

97.92

+ 8.88

97.06
95.22
71.44

- 0.86
- 1.84

77.82

4 6.38

76.44
72. 20

- 1.38
- 4.24

92

12.85
13.51
10.78
10.67
11.74
10.82
13.24
11.22
13.49
9.80
10*33
11.62
10.83
9.89
12.37
9.99
11.61
11.01
12.10
12.26
16.01
13.25
10.46
10.39
11.97
12.54
12.84
14.45
13.41
11.26
9.52
9.01
11.41
13.31
10.68
10.69
11.62
13.02
11.66
12.09
13.75
10.08

area.
: Percentage.
Gain or
Water.
loss of
carbohy­
drates.
56.78
- 2.07

53.20

+ 0.96
0.92

54.38
54.96
54.41

+ 0.25

51.20

- 3.16
+ 1.29

52*78
55.83
57.44

+ 1.54

53.93

- 0.76
- 0.60

55.41
58.09
55.74

+ 3.91

53.32

- 5.55
- 0.07

54.04
55.86
56.86

+ 0.87

53.72

+ 0.57
- 2.15

55.54
57.77
57.32

+ 1.89

53.75

- 0.73
+ 0.01

55.61
58.05
60.75

+ 0.04

57.46

+ 2.09
- 3.67

58.02
61.17

Table 27. Analysis of apple leaves* August 3 to 4, 1933. Variation
in dry matter, water content, and acid hydrolyzable carbohydrates
estimated as glucose.

carbohy­

drates.

9.86
10.31
9.62

+ 0.45
0.69

56. 33
52.88
55.61

H
CD

Time

area.
: Percentage
Gain or
Water
loss of

Cl
«

Variety

Grams per square meter leaf
Dry
Gain or
Carbohy­
matter.
loss of
drates.
total
dry
matter.

10.67
11.30
11.13

+ 0.63
- 0.17

54.72
51.92
54.51

+ 8.14
- 1.94

9.06
11.42
10.53

+ 2.36
- 0.89

56.18
52.84
54.66

72.30
81.48
78.80

+ 9.18
- 2.68

11.82
13.90
12.47

+ 2.08
- 1.43

56.12
52.90
54.61

7:00 a. m.
4:30 P* m.
7:00 a* m.

71.22
78.54
75.72

+ 7.32
+ 2.82

8.38
11.20
10.45

+ 2.82
- 0.75

58.39
56.34
58.13

Wealthy

7:00 a. m.
4:30 P* m.
7:00 a. m.

88.88
98.76
92.86

+ 9.88
- 5.90

12.03
13.37
12.38

+ 1.34
- 0.99

53.96
52.41
54.60

Livland

7:00 a. m.
4:30 P- m.
7:00 a. m.

75.72
82.22
76.20

+ 6.50
- 6.02

11.01
12.06
10.62

+ 1.05
- 1.44

59.79
57.56
59.38

7i,00 a* m.
4:30 P- m.
7;00 a. m.

77.02
86.18
81.76

Winesap

7:00 a* m.
4: 30 P- m*
7:00 a* m.

88.48
98.52
93.34

York

7:00 a. m.
4:30 P- m.
7:00 a. m.

72.46
80.60
78.66

Gano

7:00 a. m.
4:30 P- m.
7:00 a. m.

Delicious

93

+ 9.16
- 4.42
+ 10.04
I

Jonathan

Table 28. Analysis of apple leaves. August 9 to 10, 1933. Variation
in dry matter, water content, and acid hydrolyzable carbohydrates
estimated as glucose.

Variety

Time

Grams per square meter leaf
Dry
Gain or
Carbohy­
matter.
loss of
drates.
total
dry
matter

Jonathan

7; 30 a. m.
4:30 P* m.
7:30 a. m.

86.92
92.22
90.58

Winesap

7:30 a. m .
4:30 P* m.
7:30 a. m.

99.16
104.28
98.44

York

7:30 a. m.
4:30 P* m.
7:30 a. m.

Gano

area.
: Percentage
Gain or
Water
loss of
carbohy­
drates.

4 5.30
- 1.64

11.25
11.52
11.86

5.12
- 5.84

12.39
13.24
12.04

80.00
82.38
81.08

2. 38
—- 1.30

10.16
10.33
10.05

7:30 a. m .
4:30 P* m .
7:30 a. m.

80.70
87.36
84.42

4 6.66
— 2.94

10.41
11.83
10.97

Delicious

7:30 a. m.
4:30 P* m.
7:30 a. m.

77.90
80.90
78.10

3.00
— 2.80

8.83
11.09
10.42

2. 26
— 0.67

55.82
55.19
56.00

Wealthy

7:30 a. m.
4:30 P* m.
7:30 a. m.

99.53
102.68
100.00

3.12
- 2.68

14.50
14.91
14.66

0.41
— 0.25

54.08
52.46
54.77

Livland

7:30 a. m.
4:30 P* m.
7:30 a. m .

75.18
79.9 6
81.00

4 4*78
4 1.04

10.57
11.14
10.96

0.57
0.18

58.25
57.88
55.37

94

4

4

4

4

0.27
0.34

54.05
52.50
55.09

4

0.85
— 1.20

52.55
50.50
54.12

4

0.17
— 0.28

56.94
53.34
55.11

4 1.42
—■ 0.86

53.57
52.66
54.58

4
4

4

4

4

Table 29• Analysis of apple leaves* August 25 to 26, 1933. Variation
in dry matter, water content, and acid hydrolyzable carbohydrates
estimated as glucose.

Variety

Time

Grams per siquar e meter leaf
Dry
Carbohy­
Gain or
matter •
drates.
loss of
total
dry
matter.

area.
: Percentage
"Gain or
Water.
loss of
carbohy­
drates.

Jpnathan

7;30 a. m.
4:30 p. m.
7:30 a. m.

93.16
100.38
91.44

+ 7.22
- 8.94

13.66
14.62
13.14

+ 0.96
- 1.48

51.83
49.76
55.62

Winesap

7:30 a. m.
4:30 p. in.
7:30 a. m.

99.10
102.82
98.48

+ 3.72
- 4.34

14.53
16.04
13.72

+ 1.51
- 2.32

50.08
50.39
54.08

York

7:30 a. m.
4:30 p. m.
7:30 a • m*

85.20
88.32
84.10

3.12
- 4.22

13.40
13.04
12.25

- 0.36
- 0.79

52.49
51.74
53.95

Gano

7:30 a. m.
4:30 p. in.
7:30 a. m.

97.16
101.36
94.30

+ 4. 20
- 7.06

14.86
15.71
14.52

+ 0.85
- 1.19

51.73
50.25
54.00

Delicious

7:30 a. m.
4:30 p. m.
7:30 a • m.

89.34
91.24
86.92

+ 1.40
— 4.32

15.78
15.57
15.41

- 0.21
- 0.16

52.80
53.16
55.30

Wealthy

7:30 a. m.
4:30 p. m.
7:30 a. m.

109.88
115.24
105.98

+ 5.36
- 9.26

20.91
20.51
19. 61

- 0.40
- 0.90

49.50
50.11
52.72

Livland

7:30 a. m.
4:30 p. m.
7:30 a. m.

84.56
88.52
82.42

+ 3.96
6.10

14.69
15.03
13.46

+ 0.34
- 1.57

54.62
56.65
58.03

95

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Table 31* Analysis of apple leaves. May 2 and 16, 1934, Variation in total dry natter,
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Table

34* Analysis of apple leaves* July 2 and 3, 1934* Variation in total dry matter,
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Table 35* Analysis of apple leaves, July 9 and 10, 1934. Variation in total dry matter,
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total dry matter, combustible matter, water content, and acid hydrolyzable
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1334,
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Discussion and Conclusions
Based on the data presented in this paper, the following
conclusions may be drawn*
1.

Apple varieties differ in their internal structure when

the extent of the intercellular spaces in the spongy mesophyll is
considered*

The extent of these spaces was judged by measuring

tracings of projected images with a planimeter and a char tome ter*
2*

Orchard grown apple leaves possess more extensive

intercellular spaces than greenhouse grown leaves.
3.

Livland, a Russian variety, has more extensive intercellular

spaces than any other variety studied*

In 1933, the Delicious

mesophyll was the most compact, but in 19 34, the mesophyll in York
leaves was the most compact*
4.

During the abnormally dry and hot seasons of 1933 and 1934,

the stomata on outdoor foliage were seldom open after 9:00 a. m* and
frequently they were closed or nearly so by 7:00 a* m* or 8:00 a. m.
No afternoon opening of the stomata was observed.

Stomata on apple

leaves in the greenhouse remained open longer each morning than those
on outdoor foliage*
5.

Extreme difficulty was encountered in attesting to determine

whether stomata were closed or nearly closed.

It is probable that the

stomata which appeared to be closed were not sufficiently closed to
make them gas tight*

108

6*

Livland leaves had the fewest stomata per unit of leaf

area and the longest pores.
7.

Outdoor grown leaves have more stomata per unit of leaf

area than greenhouse grown leaves.

In some instances this difference

amounts to as much as 50 per cent.
8.

In January 1933, some one-year-old Livland, Jonathan,

&ano, and York trees were divided into two groups.

One group was

planted in a greenhouse where the day temperatures were usually 15
to 20 degrees C higher than in the house where the second group
was planted.

The leaves of Livland, Jonathan, and York grown in

the warmer house had a greater extent of intercellular spaces than
those in the cooler house.
(a)

The reverse was true with Oano#

Within each variety, the leaves possessing the more

extensive intercellular spaces produced the greater gain in total dry
matter per square meter leaf area between 8:30 a. m. and 5:30 p. m.
(b)

The intercellular spaces in the mesophyll in the Livland

leaves from the warn^r house were more extensive than those of the
other varieties g r o m in the same house.

The same was true comparing

Livland with the other varieties g r o m in the cooler house*
(c)

Livland leaves accumulated a greater gain of dry matter per

unit of leaf area than the other varieties within each group.
(d)

Leaves on potted trees in the greenhouse have a greater

amount of water in them than orchard grown foliage.

109

9*

In January 1934, Livland and Delicious trees were planted

12-inch, clay pots in the greenhouse.

In July 1934 these trees were

transported to the orchard so that photosynthetic Btudies of the
greenhouse grown trees could be made under the same environmental
conditions as the orchard trees.

There are marked differences in the

spongy mesophyll of the leaves of greenhouse grown trees contrasted
to orchard grown leaves.

The four groups of leaves rank as follows

in the measurements of the intercellular spaces in the mesophyll:
(1) orchard grown Livland, (2) orchard grown Delicious, (3) greenhouse
grown Livland, and (4) greenhouse grown Delicious.

The day gain

in total dry matter per unit of leaf area stands in the same relative
order.
10.

In 1933, the amount of total dry matter accumulated per

square meter of leaf area of the seven varieties used was apparently
markedly influenced by the maximum daily air temperature.

On the days

when the temperature was extremely high, the gain in dry matter was
less than on days when the temperature was lower.

In 1934, however,

the inhibitory action of high temperatures did not become manifest,
probably because of the great deficiency of soil moisture which became
a limiting factor in leaf activity.

On several days in 1934 dry matter

determinations were made when the leaves were partially wilted.

- 110 -

11*

Apparent photo synthetic determinations were made on three

consecutive days in the greenhouse in 1934 by measuring the amount of
carbon dioxide absorbed by a leaf on a potted Delicious tree and by
one on a potted Livland tree.

Dry weight increments, ash determinations,

and carbohydrate analyses were made from other leaves collected from,
these trees.
(a)

On two of the three days the Livland gained more in dry

weight per unit of leaf area than the Delicious foliage.

After being

corrected for ash content the mean gain of the dry matter for the three
days was 2.81 gms M 2 5 hrs. for Livland and 2.30 gms M 2 5 hr s. for
Delicious.

The mean accumulation of total acid hydrolyzable carbo­

hydrates estimated as glucose was 0.56 gm M 2 5 hrs. for Livland and
0.22 gm M 2 5 hrs. for Delicious.
(b)

The mean amount of carbon dioxide absorbed by the Livland

leaf was 4.81 gms M
(c)

2

5 hrs. and for Delicious 1.89 gms M

2

5 hrs.

Subsequently the two leaves used in these continuous gas

stream studies were killed, dehydrated, and imbedded in paraffin after
which mounts on slides for microscopic study were prepared.

As described

earlier, the intercellular spaces in the mesophyll were measured.

The

cross sectional area of these spaces for Livland was 107.71 cm2 per unit
area compared to 87.01 cm2 for Delicious.
(d)

The ratio of the average weight of carbon dioxide absorbed

per square meter of leaf area for the three 5-hours run to the average
weight of the total acid hydrolyzable carbohydrates accumulated for the
same area and time is 8.59 for Delicious and 8.58 for Livland.

- Ill -

12.

York leaves possess the following distinguishing

characteristics;

(a) a lower percentage of ash than Jonathan,

Delicious, or Livland, (b) usually a lower water content than Wine sap,
Jonathan, Delicious, Livland, Wealthy, or Gano, (c) a lower total
dry weight per unit of leaf area than any of the other varieties, with
the occasional exception of Gano, (d) a more compact mesophyll than
any of the other varieties studied in 1934*
13.

During both 1933 and 1934, York made lower daily gains in

total dry matter per square meter leaf

area, and lower gains in total

acid hydrolyzable carbohydrates estimated as glucose, per square
meter leaf area, than any of the other varieties included in these
studies.

The varieties used in addition to York were:for 1933, Livland,

Gano, Wealthy, Jonathan, Winesap, and Delicious; for 1934, Livland,
Jonathan, and Delicious.

It may be considered that the distinct

biennial bearing habit of York may be due in part, to the fact that the
foliage is relatively inefficient, photosynthetically, which in turn
appears to be due to son© of the items listed in the preeeeding paragraph.
14.

Livland leaves on the average have greater weight per unit

of leaf area than any of the other varieties used in 1934.

In many

instances during 1933, Wealthy leaves weighed the most per unit of leaf
area.

This is probably due to the extensive palisade development of these

two varieties.

The water content of Livland leaves is usually higher

than that of the other varieties.

These two characteristics in addition

112

to the great extent of the intercellular spaces in the mesophyll
probably account for the fact that Livland, ranked first in average
daily gains in total dry matter and total acid hydrolyzable carbohydrates
estimated as glucose per unit of leaf area,

during 1934, and that

Wealthy and Livland ranked first and second respectively in total dry
matter gains per unit of leaf area in 1933.
15.

In 1934, the ranking of the varieties

based on the extent

of the intercellular spaces and the ranking based on photosynthetic
activity as judged by the average daily gains in total dry matter, in
combustible matter, and the average daily gains in total acid hydrolyzable
carbohydrates estimated as glucose, were identical.

This ranking was

Livland, Delicious, Jonathan, and York.
16.

On six days during 1934 the total dry matter increase of

Delicious leaves on branches bearing good crops of fruit was compared
with that of leaves on branches bearing no fruit.

On five of these days

the leaves on the fruiting branches gained more in total dry matter
and combustible matter than the leaves on fruitless branches. The mean
2
gain of total dry matter for the six days was: 2.92 gms M leaf area
on fruiting branches, and 1.85 gms M 2 leaf area on fruitless branches.
The mean gains of combustible matter were 2.70 gms M

leaf area and

1.79 gms M 2 leaf area respectively.
17.

On four days during 19 34, the total dry matter increase of

Jonathan leaves on branches bearing fruit was compared with that of
leaves on fruitless branches.

On two of these days the gain was greater

for the leaves on fruiting branches and on two days the leaves on

- 113 -

fruitless branches made the greater gain*

The mean gain of total dry

matter for the four days was 2.44 gms M 2 leaf area from branches with
fruits and 1*81 gms M 2 leaf area from branches from which the fruit
had been removed*

The ash and water content of leaves near fruits

was in general higher than in leaves on fruitless branohes*

The

carbohydrate concentration was usually lower in leaves on fruiting
branches than in leaves on branches from which the fruit had been
removed*

- 114

Summary

1*

Apple varieties differ in the extent of the intercellular

spaces in the mesophyll of the foliage leaves.
2*

This anatomical difference in leaf structure is reflected

in photosynthetic activity.
3.

York leaves are the least active photosynthetically, a

characteristic which may play an important role in the distinct
biennial fruit bearing habit of this variety.

Low dry weight

per unit area, low ash content, low water content, and in 1934 the
most compact mesophyll are other characteristics of York leaves.
4.

Leaves on branches bearing fruit make greater gains in

dry matter and carbohydrates, have a higher ash content, higher water
content, and less dry weight per unit area than leaves on fruitless
branches.
5.

Leaves on fruiting branches usually have a lower concentration

of carbohydrates than leaves on branches not bearing fruit.
6.

The rate of photosynthesis was determined on 17 days in

1933 and on 20 days in 1934.

Both growing seasons were abnormally

hot and dry.

- 115 -

Acknowl e dgment s

. The writer gratefully acknowledges the help of Dr. E. C. Miller
and Professor R. J. Barnett of the Kansas State College, for their
valuable suggestions during the course of these studies and their
criticism of the manuscript.

Dr. G-. A. Pilinger of the Kansas State

College also aided in the general development of the work.

To

Dr. J. W. Crist, Professor V. R. Gardner, Professor P. C. Bradford,
and Dr. R. E. Marshall of the Michigan State College, the writer is
indebted for much helpful advice in the planning of the wo2k and
criticism of the manuscript.

116

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