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ABSTRACT

 

FOREST TREE BREEDING RESEARCH IN THREE REGIONS
OF THE UNITED STATES AND ITS APPLICABILITY
TO WEST PAKISTAN CONDITIONS

by Javaid A. Ahsan

The United States is a country with diversified
local features. The tree breeding methods adopted in the
three regions represented by Michigan State University,
East Lansing, Michigan; Institute of Forest Genetics, '
PlaCerville, California; and Institute of Forest Genetics,
Gulfport, Mississippi, to a great extent, have been
influenced by local factors.

Michigan State University has an active 10—year—
old research and teaching program. There are three major
objectives to its provenance testing work: determination
of variation patterns in conifers and hardwoods, identi—
fication of desirable ecotypes for planting in specified
areas, and at the same time providing material for further
oree’ing operations. Another set of studies is designed
to learn the inheritance of economic traits and select
00d parents for breeding, determine local variation
patterns and establish seed orchards; the objective is
being accomplished through l—parent progeny tests which

.'

are converted to seed orchards o; thinning poor individuals

 

Javaid A. Ahsan

and poor families. Allied problems, for example, deter-
mination of evolutionary and physiological factors respon-
sible for variation, juvenile-mature relationships, are
also studied. Much of the research is conducted in col-
laboration with other state experiment stations in the
north central states, under the NC—Sl project. A unique
feature of the Michigan program is the emphasis on ex—
perimental methodology, aimed at making the work more
efficient. The improvements obtained pertain to record
systems, measurement philosophy, experimental layouts,
planting methods, and nursery culture. The graduate
teaching program utilizes segments of ongoing staff pro-
jects as dissertation topics, and involves genetics,
taxonomy, and physiology.

At the Institute of Forest Genetics, Placerville,
California, provenance and progeny tests were the first
types of tree breeding studies. Fifteen years after its
establishment, in 1940, the Institute's program switched
to hybridization studies among the Various pine species
of the world. Reconnaisance of pine hybridization through
crossability studies, has remained the major line of work.
Economic potentialities of various hybrids and means to
mass produce the desirable hybrids, are also studied.

One of the major achievements of the Institute, during

the past years, is the perfection of controlled pollination

 

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Javaid A. Ahsan

techniques. In addition, some very useful hybrids have
been made.

General objectives of tree breeding research at
the Institute of Forest Genetics, Gulfport, have been
laid down as "To determine the degree of genetic control
of phenotypic characters of important southern pines and
hardwoods and to develop methods of applying this control
for the production of improved forest trees." Studies
of variation through provenance tests and progeny tests,
and also the intra- and interspecific hybridization, all
contribute to the breeding program carried out at the
Institute. A number of studies related to different
aspects of tree breeding are thus underWay. At Gulfport,
generally more sophisticated types of experimental designs,
that is, compact family block and lattice designs, are
used.

West Pakistan consists mainly of arid alluvial
plains with bushy vegetation, where irrigated plantations
of hardwood species are raised artificially; and the
mountaineous northern Himalayan region where coniferous
species grow naturally. Important species of West
Pakistan which need immediate improvement are Einpg

griffithii, P. lonaifoli , Dalberria sissoo, Acacia

 

arabica and Populus euphratica. Cooperative provenance

tests, such as those underWay at the Michigan station

 

 

Javaid A. Ahsan

with the cooperation of NC-Bl will deserve top priority,
and should involve nursery experiments, permanent field
plantations, and determination of juvenile-mature corre—
lations. These should be followed, within a few years,
by open—pollinated progeny test—seed orchards designed
to determine the variability pattern within local popu—
lations and also obtain a modest amount of genetic im—
provement of actual planting stock. These will follow
the Michigan pattern. At a much later date it will be .
desirable to follow the 2—parent progeny test designs
used at Gulfport. The pins crossability patterns worked
out at Placerville will be used to establish breeding
arboreta consisting of the species most likely to cross
with the two native pines, and pine hybridization will
for the near future be limited to production and per-

formance of hybrids involving those two species.

_,_ .‘h

 

 

FOREST TREE BREEDING RESEARCH IN THREE REGIONS
OF THE UNITED STATES AND ITS APPLICABILITY
TO WEST PAKISTAN CONDITIONS

By

Javaid A. Ahsanfl”

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE
Department of Forestry

1967

 

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ACKNOWLEDGEMENTS

I sincerely express my gratitude to Dr. J. W.
wright (Forestry Department, Michigan State University)
under whose guidance the present study was made.

I am also indebted to Drs. F. I. Righter, N. T.
Mirov, W. B. Critchfield, W. J. Libby, R. M. Echols,

E. B. Snyder, D. M. Schmitt, and O. 0. Wells, with whom r
very useful discussions were made during the period of

my study at Placerville, California and Gulfport,

Mississippi.

Thanks are also due to Le Roy Johnson, Jack
Carpender, E. F. Kimbrough, Mark Blankensop, of the
Institute of Forest Genetics, Placerville, and Jim
Hammaker, of the Institute of Forest Genetics, Gulfport,
for their help in explaining and demonstrating various
field and laboratory techniques.

Finally, I wish to thank the Agency for Inter—
national Development and the United States Department
of Agriculture, who enabled me to complete the study by

arranging my trip to California and Mississippi.

_ii..

TABLE OF CONTENTS

Chapter
I. INTRODUCTION . . . . . . . . . .

II. TREE BREEDING AT MICHIGAN
STATE UNIVERSITY . . . . . . .

III. TREE BREEDING AT PLACERVILLE,
CALIFORN-IA O C C Q C I O O O C

IV. TREE BREEDING AT GULFPORT,
MISSISSIPPI . . . . . . . . .

V. TREE BREEDING IN WEST PAKISTAN .

 

APPEDIDIX o c o a o o o o a o a o o o c o c o o o c 95
BIBLIOGRAPHY o o c o o c o a o o o o o t o a o o l 104

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CHAPTER I
INTRODUCT ION

 

CHAPTER I
INTRODUCTION

This thesis is the outcome of the author's year
long stay at Michigan State University as a student of
Dr. Jonathan wright, and a three month summer trip to
the Institute of Forest Genetics at Placerville, Cali-
fornia, and Gulfport, Mississippi, during 1966.

In a vast country like the United States there
is a wide range of climates, geographical and botanical
features, and public needs. Choice of forest tree breed—
ing approach in a particular region is governed in large
measure by local factors. The three breeding stations
involved in this study, Michigan State University,
Institute of Forest Genetics Placerville, and Institute
of Forest Genetics Gulfport, thus have different
approaches.

With the expanding field of forest genetics and
the dynamic advancement in the related sciences, there
is constant change in tree breeding methods. Therefore
in the present study I tried to learn the best methods
for present conditions as well as the processes by which

these methods change.

-2...

 

 

-5—

The three research stations have engaged in
breeding activities for many years. In order to deter-
mine how their experience can help implement tree breed—
ing in Pakistan, an appraisal was made of the techniques
employed at each center and then an attempt was made to
evaluate the adaptability of each method to conditions
prevailing in Pakistan.

Chapters II, III, and IV deal with the objectives
and methods of tree breeding followed at each of the
three research stations in the United States.

Chapter V embodies a description of the forest
conditions in West Pakistan, necessity and objectives of
tree improvement, and possible approaches to the problems
in the light of breeding principles pursued at the three

American centers of tree breeding.

  

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CHAPTER II
TREE BREEDING AT MICHIGAN
STATE UNIVERSITY

 

CHAPTER II
TREE BREEDING AT MICHIGAN STATE UNIVERSITY

The history of forest genetics at Michigan State
University is divisible into two phases, prior to 1957
and after 1957.

The earlier phase dates back to a time before
forest tree breeding was a recognized branch of science.
During the earlier years, the only aspect of forest
genetics emphasized at Michigan State University was the
introduction of exotic species. Professor W. J. Beal
and, later, Professor E. E. Bogue, hoped to find species
that would produce good wood quality and at the same
time grow as fast or faster than the native species.
Bogue started a forest nursery in 1904 from which western
pines and firs as well as eastern white, jack and red
pines and white spruce were sold for forest planting.

He also started an arboretum for his first trials at
East Lansing with the hope that progeny tests would
follow. His experiments included many pine species:

P. onderosa, P.scopulorum. Ptjeffrevi. P.flexilis,
P.rigida, and P.strobus. Other conifers such as, Pigga

bies, P.en elm nni, and Larix decidua were also grown.

-5...

 

-5-

In 1917, Professor Chittenden established a trial
plantation of Populus cangdensis and P.deltoides. Intro—
duction and trials of various varieties of poplars and
hybrids continued through the next four decades. In
the meantime, trials of exotic conifers were initiated
at the W. K. Kellogg Forest, in the early 1950's.
Ponderosa, Scotch,lodgepole, Austrian and Japanese red
pine were established in forest plantations each of one
acre or more. No tests on racial variation were estab-

lished before 1957.

 

In 1957 Prof. Jonathan Wright joined the faculty
of Michigan State University as a member of the forestry
department. Systematic efforts were started to improve
the trees of the region through the application of
genetics. The tree improvement work was speeded up
after participation of Michigan State University in the
regional NC—Bl project in 1960. NC-Bl is a cooperative
project, carried out by ten tree breeding agencies from
different North Central states, with the financial
assistance from the federal government. It is titled
”Tree Improvement through Selection and Breeding of
Forest Trees of Known Origin." An introduction of the
project with special reference to Michigan State Univer-

sity is given in Appendix I.

   

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Fgrestg 9f Michigan.—-Natural vegetation types
in Michigan are spruce—fir, beech-maple—basswood—yellow
birch—hemlock, jack pine-red pine-white pine, and aspen.
Total annual precipitation averages about 30 inches of
which 56 percent falls during the Warm months, May
through September. Irregular drought periods of 10 to
50 days, with little or no rain, are characteristic
during the growing season. The mean annual temperature
for the whole Lake States Region is about 44°F. Maxima
of 90° to lOdDF. are recorded throughout the region
almost every summer. Minima below 00F. are recorded
every winter and have gone as low as —49°F. in the upper
peninsula. Topography is typically level to gently
rolling. Soils range from dry sandy soils occupied
by jack pine or oaks, to loams With good conditions of
moisture and fertility.

Unchecked forest fires in the wake of logging
operations were an important factor responsible for the
deterioration of growth and depletion of soil fertility.
Aspen, jack pine and scrub—oak have characteristically
invaded vast areas cut-over and burned lands. The
forested area of the Lake States was 104 million acres
at the time the first white men came. This area has
been reduced to 56 million acres, of which one-fourth
is barren or not restocking. A survey of the Michigan's

forest resources reveals that natural mortality accounts

 

—8—

for more than one third of the total annual removal.
Diseases are the greatest single cause of losses. Red
pine, which is Michigan's most planted tree is attacked
heavily by the European shoot moth.

The biggest job ahead is to improve the state's
forests by reforesting the poor forest areas with the
proper tree species which can exploit site quality to
the maximum. In the following account, the role of

Michigan State University will be evaluated in this task.

 

Major Objectives of Breeding and
Lines of Work

The objectives of tree-breeding at Michigan State
University may be summarized as follows:

i. For important tree species to determine
the pattern of variation and the best
sources for planting in specified
localities; to determine the evolution-
ary and physiological factors responsible
for variation; and to furnish material
for future breeding work (this objective
is common with that of NC—5l).

ii. To learn the inheritance of economic
traits, select good parents for breeding,
determine local variation patterns and

establish seed orchards.

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The first objective is being achieved by prove-
nance studies. The preliminary results indicate the
acceptable varieties. However, in order to obtain more
concrete results, the studies are to be continued for
the next several decades. At present most of the ex-
periments are in various stages of establishment. They
must be followed because every year the information be-
comes more valuable.

Very few hardwoods have been included in the
studies of racial variation. A major hurdle has been i
the lack of satisfactory methodology which has so well
been worked out for conifers. Difficulty of storing
seeds from one year to the next (e.g. in red oak,
cottonwood, and sugar maple); difficulty of planting
and the need for intensive weed control, have been major
problems.

Work on the physiological aspects of racial
variation has started recently. It will involve chemical
and other detailed analyses of material collected in
well established outplantings. Such studies are in-
herently expensive and require specialized equipment
and personnel. These conditions were met in the past
three years.

Material for future breeding work will be fur-

nished by selected races. By crossing selected races

   

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within species and amongst different species, real im—
provement may be obtained. Nevertheless, selection of
best races and subsequent improvement based upon them

does not in any way exhaust all the possibilities of

tree improvement. A second set of objectives therefore,
envisages learning the local variation and inheritance

of desirable characters, and selecting the best individuals
for breeding and establishing seed orchards.

In order to aid the achievement of the major
objectives, research is needed on many allied problems.
One of the important problems is the long time required
in breeding work. It can be attacked by determining the
juvenile—mature correlations and by finding out the ways
to induce early flowering. Research on juvenile—mature
correlations has been started under NC—5l project in that
all experiments have been measured when one and two year
old and at intervals after the trees have been transferred

to their permanent locations.

Relationship of the Breeding Approach
pp Local Conditions

Important local factors which have led to the
choice of provenance studies as the main breeding method

in Michigan State University, are discussed briefly.

 

 

..ll—

Climate gpd edgphic factors.—-For an efficient
provenance or progeny test, climate and soil favorable
for growth are necessary. Low mortality is of particular
importance. Experience indicates that 85 percent survival
is desirable in all tests. Another requirement is that
the area should be uniform. Thus level or rolling topo-
graphy is more desirable than a hilly site. Michigan
and other areas of the North Central states, satisfy such
requirements. In the prevailing climate, with adequate

care, initial mortality can be kept below 5 percent.

Plapting practices.--Mention has already been
made of the large scale reforestation needs in the region.
A start toward this end has been made. By the end of
1948 the total area planted in the Lake States was about
1,500,000 acres. Considering all North Central states,
at the end of 1962, 5,907,000 of forest and 951,000 acres
of tree windbreak plantings had been established. During
one year, 1962, 161,000 acres of forest land and 28,000
acres of wind breaks were planted within the region.

The greatest challenge before the tree breeder of this
area is to allow the planting of only superior varieties
of both native and exotic trees, lest the mistake once
committed in planting inferior Varieties of Scotch pine
is repeated. Until improved Varieties are available for

massive planting programs, it will be worthwhile to plant

     

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the best natural races. Selection of races and individual
trees constitute this preliminary phase of tree improve—

ment in the region.

Chpistmas trees.--Michigan is second to Washington
in Christmas tree production. Provenances which satisfy
the requirements of good Christmas trees are therefore of
much value. Growers are already making use of the 8-year
data on Scotch pine and white spruce, buying seed from
regions which yield trees with acceptable fall coloration

and growth rate.

Participation of Michigap State University in
NC-51 project.—-Michigan State University is an active
member of NC—51 project whose main part is provenance
research, or testing of trees from different parts of
species' natural range. Tree breeding research in the

University should obviously be concentrated on this field.

Arboretum fgcilities.——Michigan State University
campus is itself a large arboretum. The collections,
while growing, are still deficient in the conifers. There
are smaller arboreta in nearby Battle Creek and Ann Arbor.
Interspecific hybridization will await the development of

better collections.

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Salient Featuraa of the Methgdglggz
at Migpigap State Univapsipy

In executing the tree breeding work, foremost
considerations kept in view at Michigan are: reducing
the amount of work without appreciably affecting the
accuracy of results, lowering the cost of the experiments
to the minimum, and simplification of methods. Conse-
quently, allied research is carried out to find out ways
and means to achieve these goals. Methods have been
developed and certain rules laid down to be followed at
every stage of the experiments. Distinctive features of

the methodology are given below:

Nursepx aatablishment apd lifting.——In order to
make full use of tractor driven mechanical equipment,
trees, seedlings and transplants are grown in 45—inch-
wide nursery beds which are prepared, sprayed and lifted
by machine. All mapping is done in terms of distance
rather than sequence. To re-establish the plot identities,
if nursery stakes are removed, measurement with a tape
is performed. A nursery map is always prepared before
actual sowing.

A randomized complete block design is generally
used. Seedlots are allotted to numbered packets which
are randomized within replicates and the random order

recorded on the nursery map. The randomization is done

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by shuffling seed packets rather than by tables of
random numbers.

Masking tape field labels, inscribed with a
black China-marking pencil are used. Labels are attached
close to the top of trees, with right side up, so as to
be read standing up.

A mechanical lifter is used for loosening the
trees. With the aid of a master list, markers are
placed in the aisle along each seedlot to be included in
a plantation, and seedlots are marked for pulling.
Pulling starts at the corner diagonally opposite the plot
stake which is not disturbed until the last tree is out.
Each seedlot is tied separately. Each seedlot bundle
contains some labelled trees.

Seedlings are transported to the planting shed.
Then, in each seedlot labels are added to make the total
number equal to the number of replicates. Each seedlot
is then separated into 4—tree plot bundles each of which
contains a labelled tree. Roots of seedlings in plot
bundles are dipped in water to save them from drying out
during handling. Each plot bundle is dropped into a
separate pile representing a replicate. Plot bundles
of each replicate are packed separately and labelled as

1, 2, 5 etc.

 

 

flaad control.-—For planting, uniform sites of
average to above-average quality, free from any shrub or
tree growth, are chosen. They are prepared by plowing
furrows to eradicate all the weeds well before the plant-
ing is carried out. Or weei-free strips are prepared by
spraying dalapon or amino-triazole six months ahead of
planting time. Simazine or Simazine plus amino-triazole
are used subsequent to planting. 2,4,5,—T is also used

for killing the brush.

P1apting.-—Standard planting technique is used
while planting, taking care to place root crown about 1
inch below the soil surface and to firm the soil about
the roots. No planting is done within forty feet of
existing tall timber.

Approximate replicate and plantation boundaries
are marked by temporary stakes in a manner designed to
make within replicate variation as small as possible.

In aligning the plots, care is exercised to see that they
follow obvious site gradients. All the plots are kept

in the same direction, with straight rows. Square or
approximately square replicates with regular outlines
(except one or two on the border line of the plantation)
are usually used.

A border row consisting of the same species is

planted around the entire planting after all the

 

 

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-15-

experimental trees have been planted. Identity of all
the border rows is maintained.

I Plot bundles are taken out from the numbered
replicate bundles and planting is done in a random order
within a replicate. The labelled tree is planted in the
same relative position in each plot. In order to avoid
confusion if a packer places less or more than the proper
number of trees per bundle, a space is left unfilled or
the excessive tree discarded. In case the labelled tree
is not planted in its proper position it is replanted.
If two labels are found in one plot bundle, one is torn
off.

One of the outstanding developments made at
Michigan State University in reducing the time and labor
of planting, is the perfection of machine planting
technique. It has been successfully applied in Scotch
pine and red pine. In machine planting, it is essential
that the replicates should have nearly regular boundaries,
with their long direction in the direction of tractor
travel. A base line is established with stakes at
accurate distances along one edge of the plantation.
Assume there are three replicates lying adjacent to one
another in sequence 1, 2, 5, each having eight 4-tree
plots, sixteen 4—tree plot bundles will be required to
plant up one replicate. Sixteen plot bundles for each

replicate will be placed on the tractor in the sequence

 

.17-

l, 2, 5. While the tractor travels, a walker will untie

the Q-tree bundles and hand them to the tree planter. ‘
The planter will plant the 8 plots in replicate 1, then 1
the 8 plots in replicate 2, etc. The walker will check
the replicate boundaries to insure placement of the ‘
plots in correct order. At the end of the third repli— ‘
cate, the tractor will travel in the reverse direction,

hence the replicates will be planted in the reverse

sequence, that is, 5, 2, l. Distances between adjacent

plants are maintained uniform by planter, by observing ,

the base line or the previously planted rows.

Mapping.-—In the plantation maps, a row is always
denoted by a number and a column by a letter. Only one
entry per plot is made. A missing plot is indicated by
a double dash (——) and plots of unknown identity by a
question mark (?).

Field copies of the plantation map are prepared
as the planting progresses or soon after it is completed.
Then as soon as possible, 125 permanent copies are pre-
pared by cutting mimeographed stencils. A copy of each
map is distributed to all interested individuals.

Each map (on 8 x 10-inch paper) is a complete
report, showing not only the location of the plantation
and individual plots, but also giving all the details

regarding establishment, weather at the time of planting;

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topography; soil type and soil condition; experimental
design; plot size and direction; number of varieties;

number of replicates and other pertinent information.

Measurement.—-To ensure rapidity of work and
accuracy, one or a few traits are measured at a time.
Metric traits (height, diameter, etc.) are measured to an
accuracy of l/20th of the range between the extremes.
Presence-absence traits are measured in terms of the
number of trees per plot. Non-metrical traits such as
color, are scored according to a system of numbered grades.
In all the measurement work, only the plot means are

recorded. This device saves considerable time.

Data recording.—-There is a record folder main-
tained for every plantation. It is duplicated by carbon
after the planting is over. One copy remains with the
experimenter and the other with the ground man. Either
can measure and record in his folder and transmit the
carbon copy of the record to the other. Such an arrange-
ment provides a complete and up-to-date record system in
the two folders, all the time.

A record folder comprises five parts: planta-
tion map, pages of accession record, a set of measurement
instructions, field data sheets, and empty pages on which
to record descriptions of measurements and appropriate

notes regarding operations.

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In addition to the data and other observations,
results of Various statistical analyses are placed in

the folder under appropriate characters.

Experimental desigp.-—Theory and practice of
plot size to determine a suitable design for various
experiments has been a subject of intensive investigation
at Michigan station. It was found that for greatest
efficiency 4-tree plots seem best for provenance studies
and 8 to lO-tree linear plots, for l—parent progeny tests.

A randomized complete block design has most
commonly been used in various tests. The main advantage
of this design is that pre—location of plots is not
required. Moreover, it is far simpler to analyze sta-
tistically than more sophisticated designs.

Where the mortality is lower than five percent,
instead of going into the lengthy, text-book procedure
of calculating the values for missing plots, a short-cut
is used. The mean value for the seedlot is entered and
error degrees of freedom reduced by one correspondingly.
The value of a missing plot can also be entered as the

value of same plot in other replicates.

 

-20-

Typical Studies

Being the main line of work, investigations on
provenances, constitute the largest number of studies at
the Michigan State University station. Important species
involved are: Scotch pine, ponderosa pine, European
black pine, eastern white pine, Virginia pine, red pine,
Japanese larch, white spruce, Douglas-fir, and white fir.
Two typical studies, those of Scotch and eastern white

pines, are described in detail.

Scotch pine.-—The study consists of two phases.
The nursery phase was accomplished wholly by Michigan
State University. In the plantation phase, 51 permanent
plantations were established in eight north Central states,
including Michigan. The Michigan station pooled results
and statistical analyses for measurements of all planta—
tions through 7 years of age.

The two year nursery test was reported by Wright
and Bull (1965). Seed was procured from 108 native and
15 planted stands of Scotch pine, from 21 countries of
Europe and Eurasia. Each stand collection comprised
seed from ten or more average trees from an area of
several acres. Origin data and other pertinent informa-
tion (about locality, altitude, date of collection,
area, soil, age, height, diameter, etc.) was obtained

with each collection.

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Sowing was done in sandy loam, highly fertile
soil of Bogue Nursery in southern Michigan, in early
May 1959. The nursery soil had already been treated
with a fungicide—weed seed disinfectant (methyl bromide),
the previous autumn. Sowing consisted of placement of
row stakes and seed packets, checking packet identity
against nursery map, hand planting the seeds 1/2 inch
deep, in rows exactly six inches apart, sprinkling 1/2
inch of sand over the seeds, and covering the seedbeds
with wire-screen and lath shades. A randomized complete
block design, with four replicates, was used. Each plot
representing one origin, consisted of one 46-inch row
perpendicular to the long axis of the BOO-foot seedbeds.
Trees destined for outplanting were sown in a fifth,
unmeasured replicate with each seedlot sown in a rec-
tangular plot wherein the seeds were sown broadcast.

Germination was completed in three weeks after
sowing. Watering was frequently done. A weed-free con—
dition (with the applications of Stoddard solvent) was
maintained. Mulching was done by a 1/2 inch saw dust
layer applied before winter.

Measurements were made at intervals during the
first three years. The measurements included all visi-
ble and differing traits. General rules for the measure-

ment work have already been indicated.

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Major geographical trends were found out in all
the important characteristics. Trends for some of the
traits are reproduced below:

Three-year height: Lowest, 12 cm. (northern Finland)
to the highest, 71 cm. (Belgium). In central
Europe, the trend was found reversed and
southern origins were 50 cm. tall.

Summer foliage color: Medium green (central Europe) to
dark green (northern Scandinavia and south-
eastern Europe) to blue green (western Europe).

Autumn foliage color: Very yellow (Ural mountains) to
yellow (Baltic countries) to dark green (couth—
eastern Europe) to blue green (western Europe).

Strength of root system expressed as ease of pulling at
the age of six weeks: Easy (Germany, Belgium,
Czechoslovakia) to difficult (northern Scandi—
navia, southern Europe).

Type of root system at age 5: Shallow and fibrous
(northern Scandinavia, Siberia) to long and
tap rooted (Spain).

Earliness of formation of secondary needles: Earliest
(slow growing northern progenies), latest,
early second year (progenies from Germany
southwards).

Time of first year bud set: Mid-July (northern Finland)

to early October (Spain).

 

 

    

 

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-25-

April 1961 marked the beginning of the perma-
nent phase of the Scotch pine study. wright 23. a1. t
(1966) have given a full account of the commencement of
this phase. V

Trees from the unmeasured replicate at the
Bogue Nursery, were lifted mechanically. After lifting,
every fourth tree was labelled with masking tape and
4-tree bundles were tied, from which replicate bundles
were made. Packing was done with tree roots in moist
sphagnum moss and heavy water—proof paper wrapping.
Planting material was shipped by surface carriers.

Experimental design of all the plantations was
a randomized complete block design. In the Michigan
plantations 10 replicates were made; in others the
number of replicates varied from 4 to 10 per plantation.
Except for Illinois, where two trees per plot were
planted, 4-tree plots were used. Plots were in the form
of rows. One or two rows border each plantation.
Spacing was 7 x 7 or 8 x 8 feet, except in Nebraska,
where 7 x 14 feet was maintained. From 60 to 108 dif—
ferent origins were represented in most plantations.
Mortality ranged from 5 to 80 percent; it was lowest
for varieties from Spain, southern France, and southern
Scandinavia; highest for varieties from central Europe

and the far north.

     

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After subjecting the data to the statistical
analysis, the following conclusions, with regard to the
intervarietal differences, could be drawn:

Winter foliage color: Very yellow-~Scandinavian and
Siberian varieties; Green-~varieties from Spain,
southern France, the Balkans and Asia minor;
Intermediate--central European varieties.

Needle length: Longest--central European varieties;
Shortest--south European varieties.

Winter injury: Spanish variety (iberica) was prone to
winter damage at seven out of sixteen plantations.

Height at 5, 6 and 7 years (measured at 16 sites in 5
states): Fastest——var. hgguenensis (Belgium,
north France, Western Germany); var. east
Anglia (England); var. hgrgypipa (Germany,
Czechoslovakia). Slowest-—var. lapponica
(northern Scandinavia); var. mongolica (Mongolia,
eastern Siberia) and altaica (Altai Mountains of
Siberia).

Susceptibility to insect attack: Most resistant to
European pine sawfly—-var. pgalepgis (results
from four different sites over three years) to
white-pine weevil-—var. iberica (two years

infestation of one plantation).

-25-

Eastern white pine.--Six years results on the
geographic variation of this species have been reported
by wright et. a1. (1965). Collection of seed from 51
stands within the species natural range was accomplished
in the fall of 1956. Michigan State University received
two-year-old seedlings of fifteen origins from the North-
eastern Forest EXperiment Station in the spring of 1959.
They were grown in transplant beds for one year at East
Lansing.

The 2—1 stock was planted in two permanent field
tests in the spring of 1960, in a lO-replicated, four-
tree-per-plot, randomized complete block design, with a
spacing of 8 x 8 feet. The two outplantings are located
at W. K. Kellogg Forest, Augusta, Michigan, and Fred Russ
Forest in Cass County, Southwestern Michigan. The trees
were slit planted. Cultural treatments in the Kellogg
plantation consisted of spraying with Simazine in 2-foot
strips between the rows immediately after planting.
Amitrol-T and Simazine were applied during the second
and third years. In the Russ plantation where the soil
was a dense sod on a heavy clay, mulching was done around
each tree in two foot square and the entire planting area
was kept mowed.

Analysis of data at six years revealed that
mortality had been fairly low, but had been the greatest

for the very slow growing Minnesota and Nova Scotia

 

 

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-25-

origins and the very fast growing southern Appalachian
origins. The Tennessee origin attained the greatest
height (84 cm.); the difference was significant from all
but two origins. Next fastest growing origins were
Georgia, Pennsylvania, southern Ontario, Massachusetts
and the Catskill Mountains of New York. Foliage color
differences were not significant from one another. Time
of leaf fall was found significantly different among

different sources.

Studies on physiological variation.——Ponderosa,
Virginia, eastern white, Austrian, jack, European black
and Scotch pine provenances have been under investigation
to determine the underlying physiological mechanism
behind external growth characters. Leaves or twigs from
existing provenance tests are subjected to chemical
analysis. In this connection the gas chromatograph (for
volatile terpenes) and mass spectrograph (for mineral
elements in the foliage) are used. Also, growth-chambers
are used to subjectdifferent races to known photoperiod,
temperature, and light intensity regimes.

From what has been accomplished, it is evident
that in nearly all of the 12 elements (N, K, P, Na, Ca,
Mg, Mn, Fe, Cu, B, Zn and A1), there are strong genetic
differences within Scotch and European black pines. It
has been further indicated, for ponderosa pine, that

coastal origins have much higher contents of N, P, K, Ca, ‘

    

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Mg, Mn, and B than do the hardier interior origins. In
Virginia pine, two southern origins which have suffered
repeated winter—kill, differ from all others in a number
of elements.

Scotch pine varieties with different degrees of
winter yellowing were kept at different night and day
temperatures during the spring. The results showed that
the trees changed from winter—yellow to summer-green
following a 5—day period during which the day—time tem—
peratures rose to 60°F. Night temperatures seemed unim—
portant. Light was found necessary for the change.

(White and wright, 1966.)

Half-sib progeny tests.—-Important species on
which l—parent progeny tests are being conducted are red
pine, jack pine, eastern white pine and Scotch pine.

Work on the half—sib progeny tests was started
in 1960 and a new species has been added about every
two years. From 125 to 450 progenies are included in
each experiment, which is first run through a replicated
nursery test then planted at three to six locations in
the state. The parents in some species (e.g. red pine)
have been randomly selected and in others (e.g. jack
pine) a very careful selection for the most important
traits has been made. All except the jack pine experi-

ment have been outplanted successfully with survival of

 

 

-28—

85 to 90 percent. Jack pine is still in the nursery
stage. Early height measurements indicate that most of
the variation is associated with stand rather than mother-
tree within the stand. Differences in other traits have
not been large when the seed was collected from a limited
area such as southern Michigan (U.S.F.S. 1966).

Half-sib progeny tests are aimed at collecting
heritability data regarding the important traits and sub-
sequent development of seedling seed orchards, depending
upon the magnitude of inheritance. Selective breeding
over a number of generations leading to the formation of
desired genotypes in the seed orchards, has been planned
for some species (e.g., jack pine); in others, as in red
pine, the seed orchards are intended to be from open
pollinated progeny.

The jack pine progeny test and seed orchard
project can be described in detail. The project was
started in 1965 when parental selection of 450 superior
trees was made from natural stands. Superiority of these
trees was judged on branch fineness, stem straightness,
and height growth. Special emphasis was placed, however,
on favoring trees having non—serotinous cones. Selection
intensity was such that each selected tree was the best
of 100 to 500 dominant trees growing under more or less

uniform conditions in an area of less than five acres.

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_29_

Data on parental trees was recorded on specially
forms and included the following information:
Location of stand to nearest 40 acres.

Age, average height of dominant, stocking and
soil type of parent stand.

Average stem straightness and branch size of
parent stand, recorded as, below average,
average, or above average for the county.
Superiority of selected over neighbouring
comparable trees with respect to stem straight—
ness, recorded as +50 if the 'plus tree' was
best of 50, +70, if the tree was best of 70,
and so on.

Superiority of selected to neighboring trees
with respect to height growth and branch size,

recorded in the same manner.

After the selection of individual trees about

per tree were collected in late September or

early October of 1965 and 1966. Collection was done by

felling the trees. A separate cloth bag was used for the

cones from each tree. Each bag was numbered to correspond

to the data sheet and dispatched to East Lansing, where

cones were allowed to dry for three months before

extraction and cleaning was done separately for each lot.

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Seed from the selected jack pines was sown in
the spring of 1966, with a 4-replicated randomized com-
plete block design, accompanied by a fifth block to be
used for outplanting. Measurements of height and branch
size will be made in the nursery and analyzed immediately.
The measurable stock will remain in the nursery for five
years. At a later date, the data will be used to estab-
lish juvenile-mature correlations.

Four permanent outplantings of the jack pine
progenies will be established on available sites in lower
Michigan in the spring of 1968. In later years, two
additional outplantings will be made in the Upper Penin—
sula. Sites which are slightly above average in quality
and adaptable to machine planting will be chosen. The
following specifications have been outlined for the
permanent phase of progeny tests:

Design: A randomized complete block design, each
replicate to contain one 6-tree plot of each
progeny.

Spacing: 6 x 6 feet; trees check-planted to keep the
rows straight in both directions.

Number of replicates per planting: 10

Method of planting: Machine planting with a 5-man crew.

Rate of planting: 4000 to 5000 trees per acre.

  

 

     

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-51-

Care: Intensive, including brush control, possibly
control of herbaceous weeds through chemicals,
possible fertilization and pruning. No shearing

to produce bush—type trees.

With the start of flowering the jack pine out-
plantings will be measured. Measurement will include
branch diameter, stem straightness, height, etc. Parent—
progeny relationships will be determined by subjecting
the data to analysis of variance and correlation analysis.
Other sets of measurements will be made and statistically
analyzed at five or ten year intervals.

Subsequent to each set of measurements, the
plantings will be thinned so as to remove the poorest
progenies (family selection). In the later thinnings,
the poorest trees in the best progenies will be removed
(combined family and mass selection). The genetic
quality of the stand and of the seed produced will in-
crease with each thinning. Theoretical aspect of the
genetic gain is illustrated as follows:

a. From the 1/2 sib open pollinated trees

seed with 1/2 szG will be had. The con-
stant K refers to the heritability (portion
of variance due to additive genetic factors).
The genetic gain after mass selection is

45G.

 

-32-

b. The expected genetic gain after family
selection will be equal to 1/2 Ksz +4§Ghs.
AiGhS denotes the gain from 1/2 sib family
selection.

c. The new expected gain as a result of the
combined family and mass selection would be
equal to 1/2 KziG + ALGC. znGc represents
the fraction of total gain due to combined
family and mass selection.

RemOVal of the poorest progenies will mark the
conversion of progeny tests to seed orchards. Other
cultural treatments aiming at producing abundant cones
and ease of collecting them, may include complete weed
control, fertilization, and pruning of lower branches.

The above progeny test—seed orchards would be
only a single step in a several generation improvement
program, and because of this, they are to be replaced
at the age of 20 by F2 orchards capable of producing
even better seed. F2 orchards will involve two-parent
(full sib) progenies. Theoretically, the expected gain
from the initial full—sib seed orchard is KZhG, and after
family selection, this gain will be raised to thG +
zinS. The later thinnings, which will be a combination
of family and mass selection, will raise the expected

genetic gain still further.

 

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Selective breeding will be continued into the
third and fourth generations if progress in obtaining

better seed is shown.

 

CHAPTER III
TREE BREEDING AT PLACERVILLE,
CALIFORNIA

 

CHAPTER III
TREE BREEDING AT PLACERVILLE, CALIFORNIA

Institute of Forest Genetics at Placerville,
California, is probably the foremost institute of its
kind in the world. It was established in 1925 as the
Eddy Tree Breeding Station. The present name was adopted
in 1952. In 1955, the United States Forest Service took
over the administration and now the institute is a part
of the Pacific Southwest Forest and Range Experiment
Station in cooperation with the University of California.

The institute is located at an elevation of 2700
feet, near the lower edge of the main ponderosa pine
zone in the central Sierras. Climate around the research
station is comparatively mild; maximum temperatures
rarely exceed lOOOF. Winter minimums are usually near
20°F. Precipitation is about 40 inches, nearly all of
which falls from October to May. The soil is a deep
sandy loam of a remarkably uniform character.

It was Mr. Eddy's original intention to benefit
timber-growing and lumber production throughout the
United States. Hence softwoods, rather than hardwoods,

were picked out for the principal attention. A decision

_55-

     
  

     

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Was also taken early in the life of the institute to con-
fine most of the initial efforts to the species of one
genus. Pines were chosen for intensive breeding research,

for the following reasons. First, Pinus is the most

 

Widely distributed genus of conifers in the northern
hemisphere and the United States. The genus is particu-
larly well represented on the Pacific Coast, there being
20 different species and important varieties in California
alone. Second, various species of Einug have the most
generally useful wood of any of the genera of conifers.
Pine species supply a great part of the construction
lumber for the United States and are notable timber pro—

ducers throughout the world. Finally, Pinus is adaptable

 

to a great variety of ecological conditions.
Tree-breeding at Placerville, is directed towards

the following objectives:
A reconnaisance of pine hybridization.
Evolving methods to mass produce the hybrids.
Conducting investigations, involving short-
term studies in other fields (physiology,
taxonomy, morphology, etc.) to aid the
exploratory work and facilitate the practical

application of the results.

Egploratogy investigations in pine hybridization.-—

The investigations in hybridization are aimed directly

    

     
 
  
 

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_§7_

at the improvement of pines. The objective is accomplished
in the following way. First, production of a particular
hybrid is attempted. Then, if the hybrid is obtained,
its economic potentialities are studied through perform-
ance tests in the nursery and in field plantings. In
the course of such testing, various treatments (inocula-
tion with pathogens, forced attacks by insects, etc.)
may be applied by cooperating scientists. Criteria used
in judging the economic potentialities include rapidity
of growth, adaptability to the locality where hybrid is
to be used, cold hardiness and drought resistance,
resistance to insects and diseases.

Pine hybridization reconnaissance has yielded
data on the crossability of particular species and of
hybrids, the ease with which hybrids and their progenies
may be obtained, the inheritance of characters, and the
economic potentialities of hybrids.

Initial crosses are attempted on a small scale
Without aiming at a specific location or use and without
trying to select superior phenotypes as parents (avoiding,
however, diseased and defective individuals). If sound
seeds are produced from the crossings, they are kept
separate by seed—tree and the seeds from each tree are
tested against the wind pollinated seeds from the same
tree. The results of such a procedure indicate what in
general may be expected from other crossings between the

species.

 

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As Critchfield (1965) puts it:

Once it is established that two species can be
crossed, the ease with which they cross is of
primary importance in determining how the
hybrids can be used. If two species, cross
with great difficulty, the hybrid is likely

to find its chief use as a bridge for the
transfer of characteristics between species.

If the genetic barriers between species are
weak, however, it may be possible to capitalize
on the attributes of F1 hybrid itself.

The crossability patterns being evaluated at
Placerville, when supplemented with biochemical, ana-
tomical, and morphological studies, can ultimately pro—
vide a natural system for the taxonomy of genus Pinus.
The earlier systems of inter—relationships of pines,
proposed by Shaw (1914) and Pilger (Duffield, 1952),
are based upon morphological evidence only. A classifi—
cation based upon true phylogenetic values, would un—
doubtedly facilitate the job of pine tree—breeders all
over the world.

Devising economical methods for the mass—

production of desirable hybrids.-—A first step Was made

and large trials of the most promising hybrids were

 

started in 1946. In 1948, J. W. Duffield initiated a
program for the quantity production of hybrid seeds.
There were practical difficulties. In pines, four to
ten years pass from the time of pollination to the first
preliminary judgements on growth rate and general value

under plantation conditions. Even more time is needed

 

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-59—

to decide if the hybrids will be satisfactory for timber
production. Therefore, methods for reducing the time
element have been evaluated in the institute. Is it
possible to produce male and female flowers early in the
life of a tree? This question led to grafting attempts
with young seedlings on mature trees having abundant
flowers, on the assumption that flowering of young seed—
lings might be promoted by a hormone crossing the graft.
Mirov‘s work on hard and white pines (1949) proved such
a possibility; male flowering was induced on 2—year-old
seedlings.

In order to put the hybrid pines to practical
use more quickly, F. I. Righter (1954), suggested that
they be interplanted among regular planting stock or
among the natural reproduction on cut over lands. If
the hybrid proved unsuited to the situation, it will be
crowded out; if it proves superior it will form the
mature stand of trees. However, as pointed out by
Righter, "The economic feasibility of such a course,
depends on the total increase in yield from the hybrids
in relation to the cost of seed—production carried for—
ward at compound interest." The method has been put
into practice, and interplanting tests of several species
hybrids have been established on seven of the national

forests in California (Righter and Duffield, 1950).

       
 

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Supplementary studies.--While the production of
hybrids has been the primary objective of work at Placer-
ville, several supplementary investigations have been
undertaken. A comprehensive 11-year study of the chem-
istry of pine oleoresins was completed by Mirov (1961).
The investigation shed light on the biosystematic rela-
tionships of many of the pine species. A good example
of how chemical composition of terpene may serve taxonomic
purposes, is the relationship of Pinus helepensis and
its supposed variety Pinus brutia. In the light of
chemical differences of their terpenes, these two pines
appear to be separate species. This conclusion has been
supported by independent morphological studies and by
entomological evidence.

Knowledge of terpene composition promises to be
useful in the entomological studies concerned with insect—
repellant or insect—attractive constituents of oleoresins.
Recent investigations carried out by Smith (1966) indi-
cate an association of resin vapor toxicity with the

host specificity of the insects, Dendroctonus brevicomis,

 

D. monticolae and D. jeffreyi.

Propagation of pines by cutting and grafting is
desirable in some research work. N. T. Mirov, assisted
by E. F. Kimbrough, demonstrated that not only branchlets
but even needle bundles could be used for cuttings and

for scions in grafting. Bruseof these methods, portions

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_41-

of valuable trees from other localities may be propagated,
or as many genetically identical offspring as may be
needed can be had from any particular tree.

*An investigation on seed size and seedling size
relationships sought ways to judge inherent vigor of
young trees. It was found that several factors other
than inherent capacity to grow fast affect the size of
seedling trees for several years. Differences in time
of germination and in seed size are most important.
There was often a difference of a month or two between
the germination of the earliest and the latest seeds.
When seeds are late to germinate, the young trees have
a shorter growing seas0n the first year and are, there-
fore smaller. It may take several years for the tree
to overcome this initial handicap. Schedules have been
developed for chilling pine seeds that make it possible
to germinate all seeds within three days. The stimulus
given by the food supply in large seeds may mark the
seedlings' true growth rate for two or three years. To
overcome this difficulty, all seeds for experimental use
are weighed and segregated into classes so that the
growth of trees from seed of like weight may be compared.

One of the most critical steps in the pine
hybridization program is the provision needed for long—
term storage without loss of viability. Investigations

relating to long storage of pollen have been carried out

 

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-42-

at various humidities in refrigerators and deep—freeze

chambers (Stanley at. al., 1960; Duffield and Callaham,

1959). The results indicate that pollen should be stored

at 0°C. or below and 10 percent relative humidity. Also

pine pollen kept at —2§°C. and -50°C. has germinated well.

The studies remain underway and each year pollen samples
are removed from a deep freezer and tested for viability

and fertility.

Local Conditions Affecting the
Placerville Program

To select suitable strains of Pinus onderosa, a
provenance study was initiated by Lloyd Austin and A. R.
Liddicoet in 1929. The seed was collected from South
Dakota to British Columbia and southWard to Arizona and
California. Other smaller tests were started from time
to time. In 1957, an experiment was established to test
729 individual seed trees of ponderosa pine.

In 1940, the emphasis of breeding work swung to
hybridization because (U.S.D.A., 1948): "(i) Maintaining
experimental control is difficult, and (ii) the costs of

conducting such tests may be excesSive when compared

with the benefits to be expected from this method of pine

breeding.”

Choice of hybridization as the dominant breeding

 

  

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413—

method at the Institute during the last quarter of
century can be traced back to other factors.

Pinus is the largest coniferous genus and owing to

 

the great number and diversity of the species and varied
places in which they grow naturally throughout the north—
ern hemisphere, there are almost endless possibilities
for producing new combinations of desired characters.

An arboretum is a pre—requisite for carrying out an
efficient hybridization program. By the time hybridization
was adopted as the chief breeding method at Placerville, in
1940, an arboretum was ready to facilitate the job. The
Eddy arboretum, named after the Institute's founder, con—
tains the most complete collection of pines (75 species) in
the world. In addition to this, a large number of trees in
the arboretum are raised from the nursery stock which pro-
vide testing material. Four specific purposes are achieved
with this material. They are: (l) to study early growth
of hybrids and other pedigreed seedlings, (2) to grow pedi—
greed planting stock for the Institute's plantation tests
here and elsewhere, (5) to facilitate insect and disease
studies,and(4) to proveanddemonstrate techniques and
results.

The climate at Placerville is sufficiently mild
to permit assembling large collections of the different

species of Pinus. In this climate, all but a few of the

 

most tropical of the world's species of pines can readily

be grown to seed bearing age. The average growing season

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-44-

at the research station is approximately 225 to 250 days.

There is abundant opportunity to obtain nearby
test areas in the foothills and higher reaches of the
Sierra where the minimum temperatures range from -50° to
+l6°F. Austin stated in 1957

. . . On the Atlantic Seaboard one would have

to travel about a thousand miles, from Florida to
New England, to secure as great a range in minimum
temperatures. Thus the Institute is exceptionally
well situated to assemble for hybridization pur-
poses pines of nearly all degrees of cold-hardiness,
and to develop and test strains that will combine
various desirable characteristics with considerable
cold resistance. . . .

Before establishing the Institute, its founder
James Eddy consulted Luther Burbank who had been practic—
ing hybridization in horticulture. Burbank believed that
pines could be bred as successfully as giant daisies or
potatos and therefore he encouraged Eddy to develop such
plans for the forest trees. It can thus be assumed that
from the very beginning, Eddy Was prejudiced in favor of
hybridization.

Although as early as 1927, attempts were made to
hybridize different species of pines, yet the proper
techniques had not been developed at that time and with
one exception, those early crossings failed (U.S.D.A.,
1948). With the arrival of F. I. Righter in 1940 as the

head of the Institute, a new impetus was given to the

practice of hybridization. Righter was particularly

 

 

  

 

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interested in this approach and assisted by W. C.
Cumming and A. R. Liddicoet, he perfected techniques of
controlled pollination in pines suited to Sierra Nevada
conditions. Methods to isolate ovulate flowers, extract,
test and store pollen, to pollinate the ovulate flowers,
and to test hybrids were developed during his tenure.
Through 1964, when he retired from the Institute, some
ninety hybrid combinations, including first generation
(F1), second generation (F2), back—crosses and multi-
species hybrids, had been reported.

Interest in hybridization has not dwindled with
the retirement of Righter. His absence has been compen—
sated by William Critchfield on the staff. Critchfield
has continued interest in such hybrids as Pinus attennuatt
X radiata var. bin ta; P. muricata X attenuate; P. sabiniana
X coulteri; P. sabiniana X torre ana; and P. jeffreyi X
engelmannii.

The Sierra Nevada mountains range from 200 to
10,000 feet above sea level. Such topography offers
extreme variation in edaphic and climatic factors asso-
ciated with altitude and is therefore hardly suitable
for large racial selection studies and progeny tests
which demand high degree of uniformity in the environment

for proper evaluation.

      
  

  

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-46—
Techniques of Controlled Pollingtion

Techniques of controlled pollination have been
developed at Placerville over a period of twenty years.
Improvements have resulted in (i) isolation of ovulate
flowers, (ii) collection and preparation of pollen for
use, (iii) pollination of female flowers, and (iv)
protection of cones. When properly modified, the Placer-
ville methods are applicable to other cone bearing trees
in other regions.

For the sake of convenience, seven distinct stages
of the development of ovulate strobili are arbitrarily
recognized at the Institute. They are : (l) buds small,
(2) buds large, (5) buds opening, (4) flowers partly open,
(5) flowers maximum, (6) flowers closed, and (7) cones
enlarging.

The buds are enclosed in pollination bags as soon
as they are recognizable, commonly at the "bud small"
stage, but occasionally at the "bud large" stage.

The Institute has perfected its own pollen-proof
bag. It was found that bags of materials lacking requisite
toughness were useless for Sierra Nevada conditions.
Moisture—tight bags were rejected as the vapor from tran—
spiration would condense in them and obstruct the vision
through transparent substances. Furthermore, the weight

of water that accumulates in such bags would cause the

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.47-

limbs of some species to bend down and sometimes break
off. For quite a few years, Institute's standard bag

has been a 14.5 x 8—inch bag, made up of a piece of finely
woven Army duck, with a 5 x 4-inch window made of cellu—
lose acetate (.Ol inches thick). The bag, although tough,
proved too heavy for certain species such as Pinus glabra
and P. helepensis which have relatively slender shoots.

In recent years, a "kraft paper bag" has been used. The
bag has the same dimensions as above and is similarly
provided with a 6 x 5-inch cellulose acetate window.

Kraft paper has proved satisfactory because of its light
weight and toughness. Initial cost of Kraft bags is
higher than sausage casings, however, they can be re-used.

Before re—use, pollination bags are cleaned with
a vacuum cleaner, re—lined at the neck with a 1/2 x 6—
inch cotton wand, and sealed with gummed paper. They are
then subjected to heat (80°F.) for 15 hours to kill any
pollen. Immediately prior to use, the bags are inspected
for any cracks in the paper or window.

While bagging, if there is a choice between two
or three flowers at the "bud small” stage and six or
eight flowers at the "bud large" stage, the former is
bagged. That the latter may pass the receptive stage
before pollination is possible. To pull in long branches,
a four—inch hook attached to an eight foot length of

stiff No. 6 wire is used.

 

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-43-

The number of flowers enclosed in each bag, after
tagging the branches, is recorded on a prescribed form
which is reproduced as Appendix II. The flowers that
are used as controls are also tagged. This allows a
check on the effectiveness of pollination.

Technicians at Placerville collect male catkins
when a clear liquid oozes out when pressed, that is before
perfect maturity is attained (yellowish juicy liquid is
the indication of complete ripeness of the male catkins).
At this stage a few male catkins are mature on the tree.

In the laboratory, male catkins are washed and
dried before the pollen is extracted. Extractors consist of
sterilized 12—inch x 50-inch canvas bags and funnels
which empty through a rubber tube into glass receptacles.
A four inch circle of 60 mesh screen is soldered within
each funnel just above its spout to catch coarse materials,
such as catkin particles, dried pitch, etc., which other-
wise would mix with the pollen. A sieve of the same
diameter as the funnel is held about two inches above the
funnel by metal legs that are soldered to the bottom of
the hoop. The bags have two windows. The receptacles
are glass bottles equipped with a rubber cap in which a
cross (+) is cut to permit entry of the rubber tube into
the receptacle. When the tube is forced through the cut
in the cap, the cap makes a tight fit around the tube.

When the tube is withdrawn, the cap closes automatically.

 

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The catkin-washing operation is very carefully
done. The catkins are loaded into a funnel and washed
with a stream of lukewarm water, then the funnel is
immersed in a Warm water bath. While still immersed the
catkin-containing funnel is fitted into the lower end of
the canvas extractor. Then extractor and funnel are
removed from the water and the lower end of the funnel
plugged with cotton. Then the extractor is placed to
dry in a heated room with circulating air. Pollen is
ready in l to 5 days and is drawn off through a small
rubber tube, the end of which is sterilized (with alcohol)

after each use.

Testing the pollen.——Pollen is tested for via-
bility before use, using the 'hanging drop' method
described by Righter (1959). Holes are punched in a 2-
millimeter layer of paraffin placed in the top of a
Petri dish, and in these holes the liquid culture media
is placed. A few drops of distilled water are placed in
the Petri dish to maintain humidity. The inside periphery
of the top section is coated with vaseline to provide a
seal. For preparing the culture, a sterile hypodermic
syringe equipped with a rubber bulb and 15 gauge needle
is used. A portion of each lot of pollen is withdrawn
from its container into a separate syringe and distilled

water is next drawn into the syringe and mixed with the

 

   

 
 
  

  

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-50-

pollen by shaking. To place the culture in the Petri
dish, a drop of the pollen water mixture is squeezed
into a hole. The dish is then placed in an electric"

oven, adjusted to maintain the temperature at about 26°C.

Pollingting the flowers.—-When the flowers begin
to emerge from the buds they are watched closely so that
they can be pollinated between the fourth to sixth
(preferably fifth) stages. The materials include labelled
vials of pollen, labelled hypodermic syringes of 10 cc.
capacity equipped with rubber bulbs and 16 gauge needles
whose points are protected with corks, aluminum data tags,
pertinent data, record sheets in a loose-leaf note book,
plungers for clearing needles that become plugged, wires,
a rope, pencils and a knapsack.

When flowers are ready for pollination, the
technician forces the pollen into the bag through the
needle of hypodermic syringe. The puncture is closed by
an adhesive tape. The bag is then shaken vigorously to
distribute the pollen thoroughly around the flowers.

A properly labelled 8—inch aluminum tag is
fastened to the branch a few inches below the bag.
Stapled to the tag is a strip of colored cloth to serve
as a flag. A tag applies to all cones maturing on the
stem above it. Pollinations on the same tree are often
made over a several—day period. Pollination bags are

removed after the ovulate flowers reach stage 7.

 

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Protection of geed gnd cone.—-Cloth bags are
applied to the conelets pollinated during the previous

seasons to guard against attack by squirrels and insects.
These bags also catch seed from cones opening prematurely.
Cone bagging is generally done in the spring and before

the attacking insects become active.

Hybrids Made at Placerville

Since the inception of the Institute of Forest.
Genetics, about ninety different hybrids (Fl inter-
specific crosses, back-crosses, 3—way hybrids) have been
produced. Some hybrids were easy to make. In contrast,
Pinus nigra X resinosa yielded only 7 seeds after several
years' effort. Reports by Liddicoet and Righter (1960)
and Critchfield (1965; 1966) cover most of the hybrids
produced. Although most of the hybrids are too young
to show their true capabilities, as Duffield and Snyder
(1958) point out ”Evidence from natural hybrids found
in the wild as well as from artificial hybrids, gives a
large measure of confidence in the long term performance
of hybrids.” Some of the outstanding hybrids growing at
the Eddy Arboretum are mentioned as follows.

Pinus ponderosa X 1gtifolig shows early growth

advantages over P. ponderosa for reforestation in the

 

California region (Righter and Duffield, 1951). This

 

 

  
  
 

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combination has a long, stout tap root which can be of
great value in a region of high summer temperature and
little or no summer rain fall. It outgrows P. ponderosa
in height and diameter.

Hybrids between jack pine of the Lake States and
lodgepole pine of the Rockies and Sierra Nevada, have
the good form of lodgepole pine and a growth rate almost
equal to jack pine. It should be of value in the Lake
States and in the Rocky Mountains.

Ponderosa pine of the Sierra Nevadas P. ponderosa

 

X var. scopulorum pine in the Rockies, equals ponderosa
pine in growth rate at four years and exceeds it after
the fifth year. This hybrid presumably has some of the
hardiness of the scopulorum parent. It is a prospect
for trial in the Great Basin area and eastwards.

Jeffrey X (Jeffrey X Coulter) pine is a vigorous
hybrid which is resistant to resin midge and pine repro——

duction weevil, both serious pine plantation pests. It

 

is cold—hardy throughout much of the ponderosa pine belt.
Hybrids between eastern white and Himalayan white
pines grow faster and are more resistant to white pine
blister rust than eastern white pine.
History of the knobcone X Moneterey pine hybrids
can be given in detail. Knobcone pine is a small, thin—
foliaged frost resistant tree of the Coast ranges and

Sierra Nevada foot—hills of California. Monterey pine

 

 

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is a rapidly growing, heavy-foliaged, frost-tender
species of the central California coast. The hybrids
grow almost as fast as Monterey pine but the important
characteristic is their.ability to withstand temperatures
as low as 16°F. They should be useful as a fast—growing
ornamental tree or for windbreaks and erosion control in
localities too cold for Monterey pine. This combination
was the first pine hybrid produced at the Institute of
Forest Genetics. It has been described by Stockwell and
Righter (1946) and Little and Righter (1965).

Fresh pollen was collected from native Monterey
pine and placed on receptive conelets of a native knob—
cone pine near Placerville in April, 1927. Hand—
pollinated cones were harvested in the autumn of 1928
and the hybrid seeds from these cones were sown in a
seedbed the following spring. Twenty-eight 2—year old
seedlings were planted in the Eddy Arboretum for testing.
Planting was done at 15 x 15 feet. A few seedlings of
both parental species were planted adjacent to the
hybrids.

The hybrids withstood cold which killed 5 to 5—
year old Monterey pine seedlings. At age 16 the hybrids
were 42 to 62 feet (average 52) tall, and 9 to 19
(average 15) inches in diameter. Adjacent knobcone pines
were 40 feet in height and 10 inches in diameter. Twenty

F trees measured at age 50 (1959) averaged 75 feet high

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and 17 inches in diameter.

Cones and catkins were seen on some hybrids at
age 4; at age 10 all bore cones.

In 1940, open pollinated seeds from F1 hybrids
(actually F2 seed) and knobcone seeds were planted in a
seedbed. At age 5 F2 hybrids were taller than knobcone
pine seedlings. The growth rate of some of the F2 popu—
lation compared favorably with the best Monterey pine
seedlings.

The hybrid test started in 1927 was repeated in
1947. Results were the same as before.

Field trials of F2 Monterey X knobcone hybrids
and seven other hybrids were initiated in 1950 at three
test sites (elevation 5,200—6,400 feet) in the Stanislaus-
Tuolumne Experimental Forest of California. Air tempera—
ture at the test site could vary between ~10O and 96oF.
The seedlings were planted as 1—1 stock, 25 trees per
plot. At age 10, the Monterey—knobcone hybrids had
failed at all three sites.

Fl Monterey—knobcone hybrids are being mass pro—
duced by the U. S. Forest Service for field trials in

several places in California.

 

   

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CHAPTER IV
TREE BREEDING AT GULFPORT,
MISSISSIPPI

 

 

 

CHAPTER IV
TREE BREEDING AT GULFPORT, MISSISSIPPI

The (Southern) Institute of Forest Genetics was
established at Gulfport, Mississippi, in 1954, by the
United States Forest Service. It was preceded by the
Gulfcoast Research Center of the Southern Forest Experi—
ment Station.

The Institute is advantageously located for
research on the southern pines on which it has concentra-

ted all of its early efforts. Longleaf (Pinus oalustris)

 

and loblolly (P. taeda) pines, their natural hybrid (X
P. sondereggeri), slash (P. elliottii) and shortleaf
(P. echinata) pines, and one minor species, spruce pine
(P. glabra) occur naturally in the immediate vicinity.
Sand (P. clausa), pond (P. rigida var. serotina) and
Virginia (P. virginiana) pines, occur naturally within
200 miles. Baldcypress (Taxodium distichum) and red
cedar (Juniperus virginiana), are other conifers of the
area.

here are also excellent hardwoods. Yellow—
poplar (Liriodendron tulipifera), red gum (Liguidamber

styraciflua), cottonwood (Populus deltoides), black

~56—

 

 

 

  
  
  

  
        

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tupelo (Nygsg gylvgtica), water tupelo (N, gguaticg),
black willow (Sali; nigra), black walnut (Juglags nigrg),
Sycamore (Platanus occidentglisl, and several species of
oak (Quercus sp.), are some of the valuable hardwood
species of the area.

Climate of the region is characterised by long
hot humid summers and short mild winters. July average
temperature is 85°F., January average is 55°F. Moderate
temperatures, the long growing season (280 days), a well
distributed rainfall (annually 55 inches), all contribute

to a luxuriant vegetation.

 

Soils of the Gulf Coast are red and yellow
podzolic; they have undergone severe laterization. They
range from sandy to clay. Generally, these soils are
poor for agricultural crops and good to excellent tree
growth.

The Institute administers the 4,000—acre Harrison
Experimental Forest at Saucier, and the lOOO—acre McNeill
Experimental Forest at McNeill, Mississippi. A lO-acre
arboretum is being established at the Harrison Experi-
mental Forest.

In 1961, the Institute was assigned the specific
responsibility for the Forest Service's research on the
breeding techniques of southern hardwoods. Initial

studies are devoted to cottonwood, sweet-gum, sycamore,

 

    

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-58—

and five of the economically important oaks—-cherrybark,
Nuttall, swamp white, Shumard, and willow. The pine

work continues;

Major Objectives of Breeding WOrk

The general objective of the Institute's breeding
program is to determine the degree of genetic control of
phenotypic characters of the important southern pines
and hardwoods and to develop methods of applying this
control for the production of improved forest trees.

The work falls into four phases: the study of variation,
the utilization of this variation in inheritance and
hybridization studies, development of most promising
strains of trees, and silvicultural relations of geneti—
cally improved varieties.

Improvement of characters such as vigor, form,
wood quality, and resistance to diseases, insects, and
adverse environmental factors are sought through selec—
tion, hybridization and modification, with special
emphasis on the mode of inheritance. Economically im—
portant characters receive major emphasis. Progeny
tests, provenance tests, inter— and intra-specific

hybridization, are being employed.

 

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_59_

Relationship of Breeding Work
to Local Conditions

The South is a major producer of wood for lumber
and pulp. The population of the United States and the
world is increasing and this means a greater demand for
southern wood fibre. Of the United State's annual pro-
duction of 15 billion cubic-feet of wood, the South grows
nearly half. Of the estimated goal of nearly 20 billion
cubic feet within the next fifteen to twenty years, the
Southern states are assigned the task of growing 10
billion. Tree breeders must explore all the methods to
increase wood production. There is also a demand for
better quality.

Southern forest plantations are managed more
intensively than in other parts of the United States.
Artificial planting is common and there is a great oppor—
tunity to plant new varieties. Test planting in the
South is greatly facilitated by large private tree
planting programs.

The Institute is fortunate in having a well
qualified crew of tree breeders who specialise in dif-
ferent aspects of forest genetics. his coupled with
the existence of most modern equipment and other facili-
ties, does not warrant limiting the breeding activities

to the minimum number of approaches.

 

  

  

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-60—

Pines have been the most commercially important
trees in the South and are grown mainly for sawlogs and
pulpwood. In recent years there has been a phenomenal
increase in the use of hardwoods for the pulp. Consider—
ing only one state, Mississippi, in the late 1940's,
round hardwood pulpwood averaged less than 200,000 cords
annually, or 15 percent of the total pulpwood cordage.
Production of hardwoods since 1955 has averaged about
880,000 cords annually, or 44 percent of the round pulp-
wood output. These considerations have led the tree
breeders of the South to place more emphasis on hardwood

improvement.

Work in Progress

Variation among southern pine races.--In the
South local seed is used for reforestation. The economic
returns from following this principle, which was derived
from early research (Wakeley, 1944, 1955, 1959, 1961;
Wakeley, pp, a1,, 1951) has probably more than paid for
all subsequent tree improvement research in the several
times over. However, practical problems still exist.
In case of a crop failure, how far away is it safe to
acquire seed? Are there sources better than the local

ones?

 

—61-

At present, the Institute is carrying out several
studies relating to the seed sources of southern pines,
independently or in cooperation with the other agencies.
One such study was designed to probe desirability of
using non-local seed of slash pine. The study was started
by E. B. Snyder in 1955 when seeds collected from sixty-
four 20— to 50—year old plantations were outplanted in
the Biloxi Ranger District and the MacNeill Experimental
Forest. The parent trees were of south Georgia or north-
east Florida origin. Seeds from eight local (south
Mississippi) slash pines were also sown in the experiment.
To date no significant differences have been found among
the sources in cone—maturity, cone—length, number of seeds
per cone, percent of full seeds, seed weight, wing weight,
wing color, and seed—coat appearance. However, signifi—
cant differences in survival were found in dry years,
Florida source being poorest. There were no differences
in fusiform rust resistance.

Another seed source study on loblolly pine Was
started in 1958. Seed was collected from throughout the

range of the loblolly pine (Texas, Arkansas, Louisiana,

Mississippi, Alabama, Georgia, South Carolina, North

Carolina, Virginia and Maryland).

Single—tree Inheritance.—-Individual tree Varia-

tion exists in various species and races (Dorman, 1961).

      

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A wide tree-to—tree variation in economically important
characters means considerable progress by selecting trees
with valuable traits.

Variation and heritability studies present a
broad field of investigations to the Institute of Forest
Genetics. These investigations relate to genetic diversity
in loblolly pine and sweetgum, within Mississippi. The
objectives are: to describe and compare the genetic vari—
ation associated with individual trees and stands, and to
select outstanding genotypes, on the basis of progeny per—
formance.

The approach to the first generation of testing
consists of two steps: first, a screening of many randomly
selected families (half-sib progenies) in a few extreme
environments within the region considered, and second,
the selection and testing of outstanding families in
several varied environments. A secondary objective of
the study is the comparison of the pattern of genetic
variation in two genetically unrelated but ecologically .
associated species.

Seed collection from 678 sweetgum in 159 stands
and 567 loblolly trees in 118 stands was done in 1962—
65-64. Stands were selected along 20 latitudinal tran-
sects chosen at intervals of 15 minutes. Seedlings for
all permanent outplantings were grown in Harrison Experi—

mental Forest nursery in 1965. Three replications of a

 

 

 

        
 

    

    

     

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-65-

compact family block design were used. Progenies were
scored in the nursery for germination rate, height and
other distinguishing traits. Three outplantings of all
families of each species were established spring 1966.
Compact family block design with 20 replications of the
678 gum families and the 567 .pine families were used at
each planting. One-tree plots at a spacing of 8 x 8 feet
were employed.

Evaluation will be done for the following charac-
teristics: survival at age 1, height and diameter at
ages 5, 5, and later years, phenological differences
whenever they appear, fusiform rust infection, and crown
characteristics.

Polygenic inheritance in diallel crosses of long-
1eaf pine is another important study. It utilizes the
method of diallel cross for heterozygous organisms. The
information to be derived includes the type and relative
effects of gene and environmental action, combining
ability, effects of selfing, and differences among re—
ciprocals. Thirteen longleaf pine trees, average to
superior were intercrossed in 1956—57. Experimental
design consisted of 4-tree subplots of the compact family
block type; subplots were paired to contain the cross
and its reciprocal. Whole plots were arranged in a 9 x

10 rectangular lattice of eight replications. In the

    

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~65-

compact family block design were used. Progenies were
scored in the nursery for germination rate, height and
other distinguishing traits. Three outplantings of all
families of each species were established spring 1966.
Compact family block design with 20 replications of the
678 gum families and the 567 .pine families were used at
each planting. One-tree plots at a spacing of 8 x 8 feet
were employed.

Evaluation will be done for the following charac-
teristics: survival at age 1, height and diameter at
ages 5, 5, and later years, phenological differences
whenever they appear, fusiform rust infection, and crown
characteristics.

Polygenic inheritance in diallel crosses of long—
leaf pine is another important study. It utilizes the
method of diallel cross for heterozygous organisms. The
information to be derived includes the type and relative
effects of gene and environmental action, combining
ability, effects of selfing, and differences among re—
ciprocals. Thirteen longleaf pine trees, average to
superior Were intercrossed in 1956—57. Experimental
design consisted of 4-tree subplots of the compact family
block type; subplots were paired to contain the cross
and its reciprocal. Whole plots were arranged in a 9 x

10 rectangular lattice of eight replications. In the

 

 

 

 
   

 
    

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—64—

spring of 1960, planting was made on 24 acres at the
Harrison Experimental Forest, with a stand density of 549
trees per acre.

The longleaf diallel progenies will be evaluated
for survival, height, branch angle, needle characteristics,

and oleoresin content.

Interspecific hybridization.——One problem tackled
by the Institute of Forest Genetics at Gulfport is deter-
mination of the value of interspecific hybridization as
a method of improving southern pines. Crossability
results with the four major southern pine species have
confirmed those of Placerville. Crosses involving lob-
lolly or shortleaf are relatively easy to make. Longleaf
X slash pine hybrids are also obtained easily.

Attempts to synthesize sonderegger pine (longleaf
X loblolly) have been unsuccessful at Gulfport, as in
Placerville (Critchfield, 1965). An investigation into
the crossability and incompatibility of this cross is now
underWay at Gulfport.

Hany valuable characteristics have been noticed
in southern pine interspecific hybrids, for example, rust
resistance in shortleaf X loblolly and tip-moth resistance
in shortleaf X slash hybrids. Longleaf X slash pines
survived well, bypassed the grass stage of longleaf, were

taller than longleaf pine at age 8 and.weremoderately

 

   
  

  
    

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-55-

susceptible to fusiform rust. Shortleaf hybrids were
moderately resistant to fusiform rust and superior in
height growth to pure shortleaf pine. They were inferiOr
in growth and survival to longleaf, loblolly, and slash
pines (Schmitt, 1966).

An important phase of the hybrid evaluation pro-
gram at Gulfport, is testing a series of selected indi—
viduals and races as parents and observing the progeny
at different sites. Screening tests of most possible
interspecific southern pines were undertaken in 1957 and
1959. Analysis of four plantings ranging in age from 6
to 8 years has revealed that site, cultural conditions
and individual parents had a marked effect upon hybrid
performance. These investigations also indicate that
hybrid survival is adequate and fusiform rust resistance
of three combinations (longleaf X slash, shortleaf X
loblolly, and shortleaf X slash) is exceptional (Schmitt,
1966).

A study of shortleaf geographic races X slash
pine was installed in the spring of 1964. Controlled
crosses were made between one race of slash pine and
eight races of shortleaf pine. The main object was to
learn whether fast growing shortleaf X slash pine hybrids
can be planted north of the slash pine range.

Shortleaf X slash may prove to be of practical

value where slash pine is an accepted species for planting

 

 

 

 

—66-

and in parts of Arkansas and Oklahoma and the unglaciated
parts of central states where shortleaf is the preferred
planting species. In the former it is thought that the
greater rust resistance of the hybrids gives them an
advantage over slash pine, despite their somewhat slower
growth. In the latter, it is hoped that faster growth
and better recovery from tip-moth attack will make the
hybrids superior to shortleaf (Wells, 1965). Also,

parent-progeny correlations will be calculated.

Technigues.-—One of the greatest problems in
southern test plantations is the presence of a number of
obnoxious weeds. They compete with the trees and present
difficulty at time of measurements. One reason for not
employing machine planting is that it is difficult to
follow a straight line due to the dense cover of weeds
on the ground (personal discussions with O. 0. Wells).
Simazine, Amino triazole, Dalapon and 2, 4, 5—T have
been only partially effective. Research is needed to
keep the experimental areas weed-free.

At Gulfport, there is a trend toward the use of
more sophisticated types of experimental designs, that
is, compact family block design and lattice type design.
Compact family block design differs from a randomized
complete block design in that the individual families

from a single stand are in consecutive plots (rows)

 

 

 

      

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-67-

rather than scattered throughout a replication. This
arrangement results in maximum.precision for.detection

of differences between families within stands and slightly
underestimates the differences between stands (wright,
1965). Another advantage is the relative ease with which
the data can be recorded and tabulated (Wells, 1964).

Compact family block design is appropriate where
genotypes divide themselves naturally into groups and
where the within-group differences are smaller or of more
interest than those between groups. For genotypes whose
differences are unknown or of equal size or interest (for
example, estimation of genetic gains, combining abilities,
and components of variance), Dr. Bayne Snyder at Gulfport,
prefers a lattice design. Although it is complicated he
asserts that newly developed computer programs will ease
statistical analyses and minimize time of operations at
all stages (Snyder, 1966).

A modified form of Michigan's folder system has
been in partial use at Gulfport by O. 0. Wells. A photo-
copy of the pertinent data remains with the cooperator
and the original with the experimenter. Plantation maps
do not contain information on soil, weather, spacing,
experimental design as do Michigan maps. Standard acces—
sion record forms have recently been introduced (Appendix

III).

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To keep the identity of planting material in the
nursery, paper field labels are used by Bayne Snyder.
Inscription is done with any water-proof ink. One advan-
tage of such paper made field labels as compared to the
masking tape labels of Michigan is that inscriptions can
be made in the office rather than in the field. However,
this advantage is outbalanced by the longer life of mask—
ing tape as a field label.

At Gulfport transparent sausage casings are used
for bagging female cones. Tests are underway to eValuate
the performance of painted sausage casings to obtain
cooler temperatures.

The warm humid climate of southern Mississippi
is conducive to rapid deterioration of pollen. The pollen
storage problem is acute. Presence of 10 to 14 percent
moisture in the "dry” extracted pollen is actually the
cause of rapid spoilage of pollen in the South. Snyder
(1961) has found a correlation between the abundance of
pollen saw fly larvae (Xylea sp.) and the relative humidity
of the extractor. He has recommended that the relative
humidity should be 55 percent or less.

Southern pine pollen is stored in sealed or un—
sealed containers, and wet or dry, with variable results.
Deep—freezing pine storage method at Placerville indi—
cates the advantage over ordinary storage in the refrig-

erator. This method, however, promises to be effective

     
  

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-59-

in Gulfport only after the pollen is brought down to'a
certain minimum moisture content (personal discussions
with E. B. Snyder). Unsuccessful attempts to freeze—dry
pollen of longleaf pine under lyophlized conditions (i.e.,
suspending the pollen in Seitz—filtered bovine serum,
freeze drying and sealing in vacuum) have also been made
(Hasseltine and Snyder, 1958).

To test the pollen germination, a solution of 10
percent sugar and 75 percent agar is made. It is cooled
to 50°C. and a small drop placed on a microscope slide.
Pollen is dropped by tapping it off a small loop of wire.
Slides are placed vertically in a slide rack over water
in a sealed container and examined after 60-72 hours.

This method is fast, about 60 samples per hour
can be prepared. It does not require delicate handling,
and the large drop buffers against drying during handling.
It is easy to handle microscopically. Contamination is
minimized compared to systems having many samples close

together within one container.

 

 

    
  
 
 

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CHAPTER V
TREE BREEDING IN WEST PAKISTAN

 

 

CHAPTER V
TREE BREEDING IN WEST PAKISTAN

West Pakistan is situated between 24° and 57° N.
latitudes and 610 and 750 W. longitudes in the west of
the Indian sub—continent of Asia. Its total land area
is 510,256 square miles. Two thirds of the total area
is covered by two mountain ranges, one in the north, and g
the other in the west. The northern range is the western
extremity of the great Himalayas whereas the western
range is a part of Suleman series. The rest of the
country is deep alluvial plain of the Indus river system.
West Pakistan experiences a rigorous continental
climate. It is very hot in summer and very cold in winter.
Most of the country is arid to semi—arid. The northern
mountains receive 50 to 50 inches of percipitation while
the plains and the western mountains get only 2 to 10
inches per year. Snow fall constitutes a very significant

part of the precipitation in the mountaineous region.

 

The higher mountain peaks (above 15,000 feet) are under
perpetual snow. The lower peaks and ridges, that is, up
to 2000 feet also carry thick layers of snow 6 to 9 months
in the year. Frost is experienced throughout the province
to a varying degree in the winter season.

-71-

     
  
 

 

   
 

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Due to rigorous climate and extreme aridity, the
natural vegetation of West Pakistan is mostly xerophytic
and sparse. According to Champion's classification
(Champion, 1956), the plains'vegetation is tropical thorn
forest which preponderates in thorny leguminous species
like Acacia spp. and Prosopis snp. Along the rivers the
natural vegetation is composed of Tam rix articulata,

T. dioica and Po ulus eu hratica, which are the pioneer
species on the silt deposits left annually by the rivers
after the flood recedes. Considerable parts of these
riverine areas have been converted to timber forests,
mainly of Acacia arabica, through direct seeding. Irri—
gated plantations have also been established artificially
in the plains, where sub—marginal land and irrigation
water are made available. The primary species used is
sissoo (Dalbergia sissoo) with understory of Mgpng.
Subtropical dry evergreen forest in which Olea cuspidata
and Acacia modesta are the dominant species occurs on the
low hills and the outer Himalayas, from 2,000 to 4,000
feet elevation. Subtropical pine (Pinus longifolia)
forest lies between 5,000 to 6,000 feet in the north—
western Himalayas. Moist and dry temperate forests con-
sist of coniferous species—-Pinus griffithii (Himalayan
white pine), Cedrus deodara (deodar), Abies pindrow (fir),

Picea morinda (spruce), Juniperus macropoda (cedar) and

 

 

-73...

with some'scattered hardwoods—~Quercus incana (oak) and
Acer pictum (maple). They occur between 5,000 and 10,000
feet in the Himalayas. Alpine forests are primarily
located from 10,000 feet to timber line, with Apigg

pi drew as the important timber species.

Necessitx_for Tree Improvement
in West Pakistan

West Pakistan is an extreme wood-deficit area.
Its land area classed as forests is less than 4 percent.
Even with the present low standard of living, it imports
5 million cubic feet of timber per year. Its population
increases at the rate of 2 percent per year (F.A.O., 1961).
Its economy is expanding fast and industries are growing
rapid y.

To solve the problem of wood shortage either
additional area should be brought under forests or the
productivity of existing forests must be enhanced. The
second approach is most feasible. The forests can be
made more productive by silvicultural treatments in com—
bination with genetically better planting stock

The importance of tree breeding in West Pakistan
is obvious, in View of the growing demands of new in—
dustries. Total dependance on natural regeneration is

giving way to 1arge~scale artificial plantations. In

-74-

the plains vigorous efforts are being made to raise more
irrigated plantations.

Another factor calling for the attention of tree
breeders is the gradual degeneration of the gene pool in
the natural forests of West Pakistan. The present
genetically poor stock is the result of past selective
logging of economically superior trees, leaving the
inferior ones to breed. Such a situation is most common
in the community owned coniferous forests. In establish—
ing artificial plantations a frequent practice has been
to collect seed from the inferior or isolated trees
which are prolific seed producers (Champion, 1955).

This malpractice continues today.

Previous Work and Achievements

Foresters in Indo—Pakistan subcontinent have been
slow to appreciate the importance of tree breeding. ii. G.
Champion was probably the first man who drew attention to
the subject. In 1925 he gave an account of the twisted
fibre in the wood of chir pine (Pinus longifolia) and its
possible inheritance. In 1950 he established the first
provenance test at Dehra Dun to support his observations
regarding racial differenCes in the wood of this native

pine.

 

 

 

     

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-75-

In 1929, thirty deodar (Cedrus deodapa) and two
chir pine trees of exceptional Superiority in height,
age, form and girth, were selected. A project was initie
ated in 1955 in the Punjab area to select a good race of
sissoo (Dalbergia sissoo) which should combine a good
stem form, shape, frost hardiness and a high timber
quality. In 1941 (Anonymous, 1941), progeny from selected
parent trees from the best forests of Hala Range in 4
Hyderabad was raised in the Kathar Barrage area. Possi-

bilities of improving teak (Tectona grandis) through

 

ecotypic selection were also explored.

In 1947 the Pakistan Forest Research Institute
came into being. Research in forest tree breeding re—
ceived little emphasis, however. Cytogenetical aspects
were studied briefly by M. I. Khan, who determined somatic
chromosome number in Acacia spp. and hybrids (Khan, 1951).
In 1954 the same author emphasized the urgent need of
research in tree breeding. But it was not until 1961
that concerted efforts were made by the Silvicultural
Branch of the Forest Research Institute to systematize
the tree improvement work. A thorough search for superior
germplasm was made in irrigated plantations. As a result,
140 sissoo and 27 mulberry trees were selected as 'plus
trees.‘ Progenies raised from thirty of the selected

sissoo trees are under study. In 1964, phenotypic

.'-' .- ‘an‘; ._.-

    

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-76...

selection of 26 trees of Populus euphratica was made
after a careful survey of the riverine forests in the
southeastern region of West Pakistan. Similar work was
initiated in 1965 in the northern coniferous forests.
So far 65 chir pine and 26 Himalayan white pine plus
trees have been selected (Khan, 1966).

In order to learn the techniques involved in
hybridization, attempts have been made at the following
crosses:

Pinus longifolia X P. CanarienSiS (New Zealand)

 

Pinus griffithii X P. strobus (U.S.A.)

Pinus griffithii X P. fleXiliS (U.S.A.)

Pinus griffithii X Iggpgpr§;cqlaI(U.S.A.)

Cedrus deodara X prlibanotica (Turkey)

An arboretum at Batrassi (approx. 5500 ft. eleva—
tion) was started in 1962. It contains a number of

pine species from all over the world, besides other

conifers and hardwood species.

Tree Breeding Goals

General goals of tree breeding in West Pakistan
should be to investigate the genetics of all major
species of economic importance and to increase their
gemetic potential for superior tree quality, growth and

other important traits. Research on the reproductive

 

 

 

-77..

processes and breeding techniques should supplement any
tree improvement program.

There are a number of species in West Pakistan,
both coniferous and hardwood, which are of commercial
importance. Only a few species can be chosen initially.
Choice should be based upon their economic importance
and improvement potential. The following five species

should be given first priority: Pinus riffithii, Pinus

 

lon ifolia, Dalber i sissoo, Acacia arabica, and Populus
euphratica.
Pines are given top priority because of their

great commercial importance in West Pakistan. Also Pinus

 

is the chief coniferous genus of the northern hemisphere.
Much genetic research work has been accomplished, par—
ticularly in the United States and Europe. Many of the
methods and principles perfected in other countries can
be used in Pakistan.

Pine breeding goals should be a high rate of
growth in Pinus griffithii, improvement in oleoresin
production and wood quality, and growth rate in P. longi-
reins.

Dalbereia sissoo is the major species of irri-
gated plantations and of the riverine forests of the
former Punjab area of West Pakistan and is by far the
most commonly used species all over the plains. Its

majorw1seliesj13furniture and construction. Growth rate

 

 

 

  
   
   
 

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-78-

and stem form are the main characters needing improvement.
Stem crookedness is common in irrigated plantations.

Also, sissoo is highly susceptible to defoliator
(Plecoptara reflexa) which sometimes kills trees outright.
Varieties resistant to this insect are needed. Improvement
may also be required in the wood quality of sissoo.

Acacia arabica covers extensive areas in southern
West Pakistan and is used for tannin, lumber, mine props
and fuel. It is very frost tender. As one proceeds
north, frost casualties increase; even large poles and
young trees are killed back in cold years. Increased
hardiness should be the primary improvement objective.

Populus euphratica should be included in the
breeding program on account of its prospective Value for
pulp, plywood and matches. It has an extensive ecological
range, from the hottest parts of West Pakistan plains to
the upper limits of tree growth (Troup, 1921). Other
species can be hybridized with Populus euphratica to
produce a hybrid which would embody several desirable
characteristics. Improvement should stress better
growth rate, disease resistance and stem form.

Being an underdeveloped country, still building
up its economy, Pakistan cannot afford to embark upon
gigantic and money consuming projects at the very begin—
ning of tree improvement program. Economical devices in

breeding should be preferred at all the stages of the work.

 

 

  
   

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-79-

Pakistan is very short of trained and skilled
personnel to carry out breeding work. Introducing a
complex breeding technique would therefore not solve the'
problems. _Lack of proper laboratory and equipment in the
initial stages also call for simplicity of the approach.

The necessity for large quantities of superior
wood has been emphasized. Along with long-term breeding
methods, workers in the country must apply short cuts and

depend much upon the methods which give early results.

Some Local Considerations Involved in the
Implementation of a Tree Breeding
Program in West Pakistaa

Staff and facilities.——At present there is one
U.N.D.P. (United Nations Development Project) expert, Dr.
Mirko Vidakovic of Yugoslavia, working as an expert in
genetics in the Pakistan Forest Research Institute at
Peshawar. He will remain there for the next two years.
I will join the Institute as a geneticist in April, 1967.
It is expected that the number of permanent geneticists
in the Institute will increase within the next ten years
to four. Presently there is one technician with cross

pollination experience. Forimmediate needs, four forest

 

 

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—80—

rangers will be trained in breeding work. The training
will consist of imparting philosophy of the work involved,
and laboratory and field studies, for a period of nine
months. Ten additional persons will be trained during
the next years.

Approximately 50,000 man-hours of common labor
per year will be available for the execution of tree
improvement work.

Nursery and planting sites will be available on
state-owned land. Approximately three man-months will be
required for site preparation for each large experiment.
This will include clearing of trees and shrubs, terracing
(in the hills), furrowing, and levelling.

Existing laboratory equipment consists of con—

trolled environment chambers, refrigerators, incubators,

 

and electric computers. A modern green—house will be con—
structed in the near future with the help of U.N.D.P.

funds.

Regignal cooperative project.——Before the initia—

tion of breeding studies, arrangements will be made to
have a joint, cooperative program of research, particu—
larly on provenances, with the tree breeders of Indian
Forest Research Institute, Dehra Dun, for species of com—
mon interest to Pakistan and India. The cooperation may

be on a non—governmental basis. It will involve free

 

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-81 ...

exchange of material and data between the Pakistani tree
breeders and their counterparts in India. Annual meetings
will be held to discuss the past achievements and plans
for the future. General pattern of this cooperative
project will be similar to that of NC—El. If there would
be a fair chance that such cooperation could be obtained
after a few years, provenance studies in Pakistan will be

delayed until the cooperation becomes effective.

A lO—Year Breeding Program
for West Pakistan

The projects and experiments listed below are in
the order of priority. A time schedule has been given
only for the first experiment of this program. Experience
and extent of success gained with every experiment will
govern the time for the subsequent experiment. Again,
local circumstances (availability of funds, labor and
other facilities, unprecedented occurrences) would in—

fluence the times set for the experiments.

Project 1: Provanance studies of the three im—
portant species of West Pakistan.——All the three species
picked for provenance studies, Pinus riffithii, P. longi-

folia and Dalber i sissoo, occupy extensive ranges in the

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~82-

Indo-Pakistan subcontinent. Although there is every
reason to suspect the occurence of racial variation.in
these species, little work has been done to explore such
a possibility. The primary goal will be to determine the
best ecotypes for planting in varied localities. Also,
the experiments will be designed in such a manner as to

provide breeding material in future hybridization studies.

Experiment 1A, To determine the extent and pattern
of variation in Pinus griffithii and to select the most
promising gaographic and altitudinal ecotypest——There are
indications of interest on the part of NC—Sl members in
provenance tests of Pinus griffithii. J. J. Jokela, the
chairman of that project will be approached to work out an
arrangement utilizing PL-480 funds for execution of the
experiment.

In general the study will follow the Michigan
pattern (in this regard, Scotch pine study will serve as
a model). In 1968 request will be made to district
forest officers in West Pakistan, Afghanistan, India,
Sikkim, Nepal and Bhutan, to provide seed from 10 or more
average trees per stand. Hopefully it will be possible
to obtain 100 such stand collections. Pertinent informa—
tion on locality, altitude, date of collection, area, soil,
age, height, diameter of trees will also be obtained.

The seed will be stored dry at 40° F.

   

 
  

    

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—85-

It is intended to have one centrally located
nursery site and three sites for installing the planta-
tion tests. The nursery will be established near
Abbottabad. Planting sites will be located in Kaghan,
Gallis, and Murree areas. The exact location of the
sites will be decided after a thorough survey of the
areas. It is desirable that the sites be uniform, above
average in quality, and near the main road connecting the
nursery and planting areas. All will be within 250 miles
ofyPeshawar.

In November 1969, sowing will be done. Two weeks
prior to sowing each seed lot will be given a "cold water
stratification treatment" (Rudolph, 1950). Three weeks
prior to sowing the seedbeds will be treated with the
combination fungicide—weed seed disinfectant 'Mylone'.
Fertilizer, if recommended by the soil chemist, will be
applied. Supplementary experiments may be needed to
determine chemicals more suited than 'Milon.’ I

Local practice of sowing three-feet—wide terraced
hill nurseries, will be adopted. Each plot will consist
of a single row running across the seed beds. Distance
between the successive rows will be at least 12 inches.

A randomized complete block design similar to that used
in Michigan will be adopted. That will be easier to

analyze than the lattice designs used at Gulfport.

   

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-84—

In addition to the four replicates for nursery
measurement, a fifth will be established to provide
material for outplantings; this will consist of 8 x 5—
foot plots sown at a density of 25 per square foot. Due
to the abundance of cheap labor,these plots will be
sewn in rows, as at Placerville. Seeds will be sown
half inches deep and covered with one third of an inch
of sand. Mulching will be done by covering the beds with
polyethylene film and then burlap until germination starts
in the spring. About 50 percent high shade will be pref-
erable until August of the first year. Seed beds will
be covered with wire screen which will be removed after
germination. Damping off, if it appears, will be con—
trolled by "Captan." Control of white grubs will be
exercised by the application of Dieldrin (at the rate of
1.5 pounds per acre); of cut worms by poison bait or
D.D.T.

Adequate watering is extremely important. Fre-
quency of watering will depend upon the site. Frequent
shallower Watering will be substituted by less frequent,
deeper watering as the plants grow older.

Preparation of sites for outplanting will be
completed by 1971. All the trees, scrubs and grasses
will be cleared. Before planting a low volatile ester

of 2,4,5—T, 5 parts concentrate in 100 parts of diesel

-_“-’.'s

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9: fiiw arofq sanifi,aadh1 qneflo 16“ _
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—85-

oil will be applied as a stump spray (White, 1965). In
March of that year seedlings will be lifted for out-
planting. The outplanting procedure and design will

in general follow those used in Michigan for hand-
established plantations. Planting stock will be shipped
in a state truck driven by one of the professional genet-
icists, who can then stay at the planting site to super-
vise.

Planting will be carried out with open rooted
stock, after pruning the roots and dipping them in liquid
mud. After planting, Simazine (4 lbs. per acre in 100
gallons of water) will be sprayed over the newly planted
trees. Application will be in coarse, low pressure
(20—50 psi) water spray in 24—inch bands (White, 1965).

Due to uneven topography, it may not be possible
to locate all replicates at the same place. They can
be separated if the site conditions within a replicate
remain as uniform as possible. The trees will be spaced
8 x 8 feet. Differences in visible traits will be
measured in the nursery. Particular notice will be taken
of mortality, growth, height, stem form, disease and
frost resistance.

In order to determine whether the visible dif-
ferences between the extreme varieties are due to chance

or otherwise, the data will be subjected to analysis of

~86-

variance. Plot means will be used as items. Statisti—
cians will be consulted to carry out detailed analyses.

All failures in the test plantings will be
replaced in November 1972 with the onset of winter rains.
No further replacement will be carried out.

The nursery experiment will probably be abandoned
after 5 years, perhaps earlier. The outplantings will be
maintained indefinitely although recent experience at
the Michigan station indicates that they may yield 90
percent of their useful information by age 10. Probably
detailed measurement will be made in the permanent out-
plantings at ages 5 and 10. Thereafter just enough
measurements will be made to determine whether earlier

conclusions remain true.

Experiment 1 B. Provenance test of Pinus longi-

folia.——The extent and pattern of variation in Pinus

 

longifolia will be studied in a provenance test similar
to that already outlined for Pinus griffithii. The
nursery study will be located near Abbottabad. Permanent
plantings will be established in Murree, Haripur and

Siran forest divisions.

Experiment 1 C. Provenance test of Dalbergia
sissoo.——The extent and pattern of variation in Dal—

bergia sissoo will be studied in a similar rovenance
——£L——————h—— p

 

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—87-

test. An existing nursery maintained by the Silviculture
Research Division, Northern Zone, near Lahore, will be
utilized for the nursery test. The planting sites, all
located within 250 miles of the nursery site, are:
Machu (Leiah), Shorkot (Multan), Chhanga Manga (Lahore),
PeshaWar.

Sowing will take place in March. Planting will
be carried out with one year old sissoo stumps consisting

of 5 inches of stem and 9 inches of root portion.

Pro'ect 2: Stu of local variation in Pinus

griffithii, P. longifolia, Dalbergia sissoo and Acacia

 

arabica; establishment of seed orchards.——Experience
indicates a good deal of local variation in these species.
Phenotypic selection of superior trees in the local
populations is being done intensively. Investigations
to know the extent of local variation and heritability
of useful characters, are necessary to find out the true
value of phenotypic selections. Seed orchards are
needed to produce improved seed for implementing the
large scale planting programs in West Pakistan. The
goals are to determine the best genotypes and the extent
of heritability; to produce seed of a high genetic

quality in the seed orchards.

-88 ..

Expariment 2A. Qpen-pollinated, progapy test—
saad orchapda of Pinus griffithii.-—The jack pine progeny
test—seed-orchard project at Michigan station, will serve’
as a model.

Seed will be collected from about 400 selected
trees by felling. Details of nursery treatment and
planting given for experiment lA will be followed. A
4-replicated randomized complete block design will be
used in the Abbottabad nursery; a fifth replicate will
be used for outplanting. Measurements will be made in
the nursery intensively for five years and later, data
will be used to establish juvenile-mature correlations.

Sites for the three permanent outplantings will
be located in Murree, Kaghan and Gallies. Design of the
plantings will be a randomized complete block; with 10
replicates, and 6-tree plots. Plantations will be
measured at five—year intervals and parent—progeny rela—
tionships will be worked out. After each measurement,
the plantings will be thinned to the best individuals

in the best families.

Experiment 2B. Open pollinated, progeny test
seed orchards of Pinus longifolia.—-Details will be the
same as for experiment 2A. A nursery will be located
at Abbottabad. Plantings will be established in Murree,

Haripur and Siran forest divisions..

     
   
  

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-89—

Exparimant 2C, Qpap pollinatad prggapy taat—aaad
_.--Major details will be the

 

same as for experiment 2A. The number of selected
parents will be 60. A nursery maintained by the Silvi-
culture Research Division, Southern Zone, at Miani
research station, will be utilized for the nursery test
and raising the planting stock. Sowing will be done in
early February. A pretreatment consisting of soaking
the seed in boiling water for 20—50 seconds and then
allowing the water to cool for 24 hours, will be given.
Sowing will be done from the seed of the same year.
Permanent plantings will be located at three
above—average sites in the Hyderabad region. Planting

will be done with balled two-year old stock.

Project 5: Hybridization of Populus euphratica

 

with other species.-—Hybridization in genus Populus has
been an effective method of improvement. There are

good chances that Populus euphratica will combine with
many exotic poplars to produce progeny with hybrid vigor
and other desirable traits. The primary goals are to
produce hybrids combining ecological adaptablity of the
native Populus euphratica with good stem form of many of
the exotic species; to produce the effects of hybrid

vigor.

-90-

Experiment 5;, Populus euphratica X P. deltoides
_ hybridization.--U.S.F.S. will be requested to send pollen

from four selected trees of Populus deltoides; it will be
stored in the refrigerator upon its arrival. In late
December, about two weeks before the anthesis occurs in

P. eu hr tica, 2 to 5 feet branches bearing female flowers
will be shot down from the twenty selected individuals,

in the Hyberabad region. They will be brought to moist
atmosphere of the green house and grafted onto rooted
stocks. Pollinations will be done in late January.

From four male parents and twenty female parents,
eighty combinations will be obtained. Seed will be kept
separate for each combination. Progenies kept separate
by families will be raised in the experimental nursery at
Peshawar. Individuals from different progenies will be
compared for height growth. In order to evaluate the
advisability of juvenile selection of hybrid progenies,
25 largest and 25 smallest trees from every progeny will
be planted in a randomized block-split plot design.

The project will be continued after the first
phase and systematic hybridization experiments with the
important species will be carried out. Tests for the
effectiveness of clonal selection will be a major part

of the project.

    

  

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-91-

Project 4, Determining effective pollination
techniques in Acacia arabica apd possibilities of inter—
specific hybridization.—-Some of the exotic Acacias are
known for their hardiness and high yield of tannin. By
hybridization it is possible to combine these traits

with the site adaptability of the native Acacia.

E eriment 4 Pollin tion techni ues for Ac cia
apabica.—~In Acacia emasculation damages the tender buds
so much so that they soon drop. Hot weather coupled with
wind and storms also take heavy toll of the flowers.
Emasculation and pollination under varied conditions of
green house will be made to find a solution to these
problems.

Future hybridization experiments will depend upon
the perfection of pollination techniques in Acacia. If
successful techniques can be evolved, the following crosses
will be made during the next phase of the breeding program:

For frost resistance: A. Arabica X A. modesta,

A. jacguemontii and A, catachu.

For drought resistance: A. Arabica X A. modesta,

A. farnesiana, A. sene l, and A. jacguemontii.

For higher yield of tannin: A. arabica X A. modesta.

Project 5, Pollination techniques in Dalbergia
sissoo and intra- and interspecific hybridization.-~

‘

 

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-92-

Development of pollination techniques is necessary for
testing 2-parent progenies and combining desirable traits
in intra- and interspecific hybrids. No hybridization
has been attempted on sissoo. Due to self pollination,
sissoo trees occur in 'pure lines.’ There is thus a
chance of producing hybrid vigor in the progenies by

effecting cross pollination.

Experiment 5A,Ibllination techniques for
Dalbergia sissoo.-—During early April before natural
pollination occurs, 50 flowers will be opened and emascu—
lated. There will be no need of isolation by pollination
bags because of the self—pollinating habit. There will
be three pollination treatments: (1) rubbing split
anthers directly over the stigma (lO flowers), (2)
brushing preserved pollen over the stigma (lO flowers),
and (5) none.

In developing a pollen forcing procedure, branches
bearing flowers will be collected two weeks before
anthesis occurs. They will be brought to the green house
and pistils will be removed. Two treatments will be
attempted: (l) grafting onto the rootstocks and (2)
keeping the lower ends in the water.

Future hybridization experiments will depend upon
perfection of pollination techniques in sissoo. If suc—

cessful techniques can be evolved, the following

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-95-

interspecific crosses will be made later: Dalbergia

sissoo X D. l tif lia, and D. siaaoo X D. cochin—chinensis.

Future breeding work.——The above program will be
only a first step for the tree improvement in West
Pakistan. After this preliminary stage is successfully
completed, it can be expected that information regarding
the provenances, individual tree variation, heritability
and cross pollination in respect to particular species
will enable us to commence advanced breeding work. During
the next phase, 2-parent progeny tests designed on the
Gulfport pattern will be used for acquiring more knowledge
of inheritance. Inter—racial crosses will be attempted
to combine different desirable traits of ecotypes. Intra-
specific crosses will also be made.

Interspecific hybridization of pines will be con-
fined to the production and performance of hybrids in-
volving the two native species only. The pine crossability
pattern worked out at Placerville will be used to establish
breeding arboreta consisting of the species most likely to
cross with the two native pines. Pronounced hybrid vigor
has been demonstrated in the following crosses in the
United States (Wright, 1962): Pinus griffithii X P. strobus,
Pinus griffithii X P. monticola, and Pinus flexilis X
P. griffithii. A successful cross between Pinus longi—
Eplga and P. canariensis has also been reported (Liddicoet

 

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the arboreta.

 

 

APPENDIX

 

 

APPENDIX I
NC—5l PROJECT

NC-5l project was started in 1960 with the sup-
port of federal funds allocated under the Hatch Act, 1886.
There are 10 state agricultural experiment stations of
the north central states as well as the Rocky Mountain
and North Central Forest Experiment Stations, of the
United States Forest Service, involved.

The project is an excellent example of cooperation
amongst various agencies in the field of forest genetics.
The representatives from the above—mentioned agencies
meet once a year, discuss the progress of work and decide
about the future goals and plans. The rigidity of rules
is observed in the matters of technical cooperation.
Nevertheless, various members are free to publish the
findings independently.

In its first phase, that is 1960—1965, NC-51
project consisted of a series of provenance studies which
aimed at providing information on the broad genetic varia-
tion patterns in the major forest tree species. Range
wide tests were initiated on 19 species; tests organised

by other agencies on 6 additional species were also

~96-

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participated by several of the NC—Bl states. Important
species under investigations have been red pine, eastern
white pine, Scotch pine, ponderosa pine, Japanese larch,
balsam fir, white fir, Douglas-fir, eastern cottonwood
and red oak. The experiments are in various stages of
establishment and are continued in the next phase. The
overall objectives have been extended from provenance
tests to other aspects of forest genetics. A summary of
these objectives is reproduced below:

(a) Determine the range and pattern of genetic
diversity within selected forest tree species

(b) Determine the physiological and evolution—
ary mechanisms responsible for growth and morphological
differences

(0) Develop useful hybrids within and between
species by crossing selected races, utilizing material
in existing provenance tests as parents. Where possible
develop methods of stimulating early fruit production to
facilitate the breeding

(d) Establish the usefulness of juvenile per—
formance data and shorten testing periods by means of
juvenile—mature correlations and theoretical analysis.

Regional cooperation in the field of forest
genetics as envisaged by NC-Bl project, is necessitated

due to several factors; listed by Wright (1965), they are:

  
 
 
   
    

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First, all species designated for improvement
are being'planted in several states. Second,
the amount of improvement needed is too great
to be handled by a single agency. Third,
little of this type of research has yet developed
to the point where the most important answers are
applicable to specific localities. Most overlap
state lines. Fourth, a coordinated approach has
proved fruitful in this and other regions.

Value of NC—El to Michigan State Universipy

Michigan State University Forest-Genetics branch
is a small organization with too limited resources to
carry out large—scale experiments. Its participation
in NC-Bl project has made it possible to embark upon
gigantic studies, a classical example of which is Scotch
pine provenance test. Without the cooperation of other
states, perhaps, it would not have been possible to test
the species over a varied number of sites and gather a
wealth of extremely useful information with regard to
the geographic pattern and performance of different seed
origins.

Besides the material benefits, several useful
technical lessons have been learned as a result of a
joint approach. For example, desirability of having a
rangewide coverage,.relatively intense sampling and
field tests over a wide variety of sites, has been

realized.

 

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A previous practice in NC—Bl experiments had been V
to raise the stock to plantable size and then distribute
it for immediate planting. There were timing difficulties
and seedlings which did well under some conditions did
not perform well under others. Lining out of the stock
for one year prior to outplanting has lead to far better
results.

It has been further learned that, to find out the
details of variability pattern, 90 to 95 percent survival
and a start of all the species under weed free and uniform
conditions is a must.

Small plots, many replicated designs have proved
to be more than satisfactory in a project like the NC-Bl,

where too many origins and too many localities are involved.

Contribution of Michigan State University to NC—il
Michigan State University has all along been a
very active member of the NC-Sl project. Besides the fact
that measurementsand analyses for many of the species have
been accomplished at this station, Michigan State Univer—
sity has distributed planting stock of a number of species
to other NC-Bl agencies for the purpose of testing and
pooling down the results. Michigan has been the iniating
agency for the provenance studies of the following species:
red pine, eastern white pine, Scotch pine, European black

pine, ponderosa pine, limber pine, Japanese larch. In

  

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i
improving the methodology of experiments, Michigan has

formulated some useful rules for the measurement and

record keeping which have benefitted other members of

the project as well. Another useful device perfected at

Michigan has been that of machine planting.

     

    
 

  

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APPENDIX II

Pollination Record Form Used at the
Institute of Forest Genetics,
Placerville

 

 

 

 

 

 

l9.___.
Experiment number . Seed parent
Flowers bagged Maturity Disbagged
Cones bagged Collected
Pollination data Crop

H
3 BE? Pollen parent, Pollinations g
E m with year of m
E 53% pollen collection o . o *5

9.4 p Maturity m o m m
m 8&4 m mtm (DH
m s m a: s w sra
E4 Z o m o o

c>p o o

 

 

 

 

 

 

 

 

 

 

 

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Accession'number

////z////////

APPENDIX III

SOUTHERN FOREST EXPERIMENT STATION
LIVING PLANT MATERIAL ACCESSION RECORD

Type of material (seed, pollen, ramet, if other, specify)

 

Species or hybrid
Pedigree (if applicable)

 

Supplier's name and address (if plant material has been
sent to Southern Station by outside address)

 

Supplier's identification of plant material
Amount of material collected

Source: Lat. Long. State County Range

 

 

 

 

 

Township Section Elevation Land grant

 

Detailed directions:

 

Aspect Slope Soil

 

Physiography (floodplain, upland, ridge, etc.)
Planted or natural stand (seed origin, if planted)
Stand history
Size of collection area

Number of plants collected from
Estimate of average age

 

Associated species

 

Comments
Collected by Collection date
Intended use (study no. or other

 

 

 

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This form should be filled out whenever plant material of
any type is collected or received. If the material is to
be used in a specific study original and 1 copy are re—
quired. If a specific study is not involved the original
only will suffice. A Southern Station Accession Number
should be assigned at this time. This number consists

of 2 letters and 10 numerals as follows:

1. A 2—letter, 4-digit designation of the
Southern Station Project that is collecting
or receiving the plant material ($01801,
SOl40l, etc.). This part of the number is
already on the form.

2. A 2—digit designation year of collection
or receipt (66, 67, etc.).

5. A 4—digit number within a group of number
assigned to the collector by the project
leader. Within this group, the numbers
may be consecutive or they may conform to
some special scheme called for by a par—
ticular study.

Example: /S/O/l/4/O/l/6/6/2/O/2/l

This was the twenty—first collection made in
1966 by the man assigned accession numbers
2000—2999; Institute of Forest Genetics,
Southern Forest Experiment Station.

ALWAYS FILL IN ALL 12 SPACES.

All different collections from a given parent or ortet
should be assigned separate lot numbers.

Outgoing shipments of the plant material recorded on this
form should be accompanied by a copy of this form.

Filing.-—If a collection is made as part of a
study, the original should be kept in a central
project file and the copy with the other records
for that particular study. If the collection
is made not as part of a study the original
should be kept in the central roject file (no
copy is necessary in this case). The central
project file should categorize the accession
forms by:

I. Type of material (seed, pollen, ramet,
or other)

A. Year of collection
1. Species

a. Increasing order of accession
number

 

   

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BIBLIOGRAPHY

 

LITURATURE CITED

Anonymous.
1941. Selective breeding of babul. Ann. Admin.
Report Sind. 1940—41.

Austin, L.
1957. Ultimate goals in forest genetics.
(Unpublished).

Champion, H. G.
1925. Contribution tOWards a knowledge of twisted
fibre in trees. Ind. For. Rec. V. 11 (part 2).
70 PP-

0
——-——~—_—

1955. The importance of the ori in of seed in
forestry. Ind. For. Rec. V. 17 part 5). 76 pp.

1936. A preliminary survey of the forest types
of India and Burma. Ind. For. Rec. V. 1 (part 1).
286 pp.

Critchfield, W. B.
1965. Hybridization of the southern pines in
California. For. Gen. Workshop Proc. 1962:40—48.
South. For. Tree Impr. Comm. Publ. 22.

1966. Crossability and relationships of the
California big—cone pines. Sec. Gen. Workshop
S.A.F. and Seventh Lake States Forest Tree Impr.
Conf. Joint Proc. 1965:56—44. U.S.F.S. Res.
Paper NC—6.

Dorman, K. W.
1961. Selection as a method of Tree Breeding.
Sixth South. Conf. Forest Tree Improv. Proc.
1961:65—72.

Duffield, J. W.
1952. Relationships and species hybridization in
the genus Pinus. Zeitschr. f. Forstgenetik u.
Forstpflanzenzficht. 1:93—97.

 

-105-

 

    

 

rdsf

 

 

o .L‘-‘ ""2”“?1: - ELL-ta d EOYIBE

 

.-
I,
u 0"!
.. ._ [.7 (1.5...
" . '. .' -.L' .‘-...
.... . ._.
, . 2,1'...._‘
y
s.'. In
n.‘
a' '\ ~
. .'.
o 5 .' -.. ..
,-
e
o
x
u
. r. n
s
u
l
a
. v
c
u
a
. .. .

nerd svi?selsfi_ .nfél ' ,

. urn-BUS, If '-

 

..-u - -..-~-

 

-lO6-

., and R. Z. Callahan.
$933.24Deep freezing pine pollen. SiIVae Genetica

 

., and E. B. Snyder.
1958. Benefits from hybridizing American forest
tree species. J. For. 56:809-815.

 

F.A.O.
1961. Timber trends and prospects in the Asia
Pacific Region. F. A. O. l 6

‘ Hesseltine, C. W., and E. B. Snyder.

1958. Attempts to freeze-dry pine pollen for
prolonged storage. Bul. Torrey Bot. Club 85:
154—155.

Khan, M. A.
1966. An intloduction to research in tree breeding
in West Pakistan (Unpublished).

Khan, M. I.
1951. Study of somatic chromosomes in some Acacia
species and hybrids. Pak. J. For. 1:526—541.

1954. Forest tree breeding. Pak. J. For.
4:252—260.

Liddicoet, A. R., and F. I. Righter.
1960. Trees of Eddy Arboretum. U.S.F.S. Pacific
Southwest Forest and Range Expt. Sta. Misc.
Paper 45. 41 pp.

Little, E. L., and F. I. Righter.
1965. Botanical descriptions of forty artificial
pine hybrids. U.S.F.S. Tech. Bul. 1545. 47 pp.

Mirov, N. T.
1949. Tested methods of grafting pines. J. For.

58:768—777-

1961. Composition of gum turpentines of pines.
U.S.F.S. Tech. Bul. 1259. I58 pp.

Righter, F. I.
959. A simple method of making germination tests
of pine pollen. J. For. 57:574—576.

 

 

 

..' ens-sci :msfiamu fig
..' , - ' 7-.

.131 Jr

* , h

a“ '»

v
.. I. .
‘ l ' 0- . I
. -.. --
.1.
_ .-. r. _q
._.I- . .. : |.-3-'--..
.
- . -.
I
_l
I 9 n : '
. , |
u
I: - ll
g .
.. . .. . . .
. . . . . - 0
v I ‘
, .
fi _ . .
.
, . ., < . . . ~
,, ..
1 n .
,
4
-. . . , . . . .
.
u . . ~

 

 

-107-

1954. Forest tree improvement research in
California. J. For. 52: 681—685.

., and J. W. Duffield.

1950. Forest Genetics—~an expanding field of
research. U.S.F.S. Calif. Forest and Range
Expt. Sta. 15 pp.

., and J. W. Duffield.
1951. Hybrids between ponderosa and Apache pine.
J. For. 49:545-549.

 

Rudolph, P. O.
1950. Cold soaking-—a short cut substitute for
stratification? J. For. 48:51-52.

Schmitt, D. M.
1966. Interspecific hybridization for the
southern pines. Problem Analysis FS—SO-1401—6.
Inst. Forest Gen. Gulfport, Miss. South. Forest
Expt. Sta. 27 pp. (Unpublished).

Shaw, G. R.
1914. The genus Pinus. Arn. Arb. Publ. No. 5.
96 PP-

Smith, R. H.
1966. Resin quality as a factor in the resistance
of pines to bark beetles. N.A.T.O. and N.S.F.
Symposium. Proc. 1964:189—196. Pergamon Press.

 

Snyder, E. B.
1961. Extracting, processing, and storing southern
pine pollen. U.S. F.S. SOuth. Forest Expt. Sta.
Occas. Paper 191. 14 pp.

 

1966. Lattice and compact family block designs
in forest genetics. Sec. Gen. Workshop S.A.F.
and seventh Lake States Forest Tree Impr. Conf.
Joint Proc. 1965:12—17. U.S.F.S. Res. Paper
NC-6.

Stanley, R. G., J. Petersen, and N. T. Mirov.
1960. Viability of pine pollen stored 15 years.
U. S. F. S. Pacific Southwest Forest and Range Expt.
Sta. Res. Note 175, 5 pp.

 

1: '.'-. - ' ~ “53:3: za-w ‘t-

'_-.n aim .g‘pxbm the
an": the :J‘se’xo’i

. u
o
u I " ‘
‘ . l 1 o -
:-
_ 1
.- . -
..y
a u '
1
- . . . .
, a

..

 

-108-

Stockwell, Palmer, and F. I. Righter.
1946. Pinus: the fertile species hybrid between
knobcone and Monterey pines. Madrono 8:157-160.

Troup, R. S.
1921. The silviculture of Indian trees.
' Clarendon Press, Oxford. 5: 957— 965.

U.SIDIA.
1948. Tree breeding at the Institute of Forest
Genetics. U.S.F.S. Misc. Publ. No. 659. 15 pp.

1966. Forest tree improvement research in the
Lake States. 1965:4-6. Ed.: P.O. Rudolph.
U.S.F.S. Res. Paper DIG-lo

Wakeley, P. C.
1 44. Geographic source of loblolly pine seed.
J. For. 42:25—52.

1955. The South establishes a major pine geo-
graphic seed source study. Sec. South. Forest
Tree Improv. Conf. Proc. 1955. 6 pp.

1959. Five—year results of the Southwide pine
seed source study. Fifth South. Forest Tree
Improv. Conf. Proc. 1959:5—11.

 

1961. Results of the Southwide pine seed source
study through 1960—61. Sixth South. Forest Tree
Improv. Conf. Proc. 1961:10—24.

., D. A. Anderson, T. E. Bercaw, and others.
1951. Standardized working plan for local tests
of seed source. U.S.F.S. South. Forest Expt.
Sta. 11 pp.

Wells, 0. o.
1965. Cooperative study of shortleaf geographic
races X slash pine. Study Plan FS—SO—1401
(IG—12—6 . Inst. Forest Gen. Gulfport, Miss.
South. Forest Expt. Sta. 20 pp. (Unpublished).

 

 
     
 

       

.fidflpé

.'.E‘es'xr‘ 96-215? ..SW-EL

 
 

   

     

. -_.
1 L' i. ". 1 - I
r " i ..+ 390
-1 ' . -’.. 5: ..'. . ..' ..I-.'.Iu-fi
-' o _-..-
. -.
- " - 1' ..'... L- .. “l"
I - .. I u - e . 5 " - _. - u --
. - '.x.
e. ' J u .
.
u . .3 - "1.: Fifi-'5“
II I -
l -' '1 ' 3“ -L
a —
q . ' v I ' l .
' I- I—IIIUI-
1
" I U _-
- n l I C
.-
n
, . a r o
.
a
a
, r
. . .
s
v . I .
~ .
u - r . 4 .
1
. a
. .
. . -.
o
., . .
-

 

White, D

Wright,

-109-

1964. Geographic variation in ponderosa pine.
Silvae Genetica 15:89—124.

. P.

1965. Fertilization and weed control on Christmas
tree farms. Ext. Bul. 505. Mich. State Univ.

8 pp.

., and J. W. Wright.
1966. Pigment changes in Scotch pine induced
by controlled environments. (In Press).

J. W.

1962. Genetics of forest tree improvement. FAO
Forestry and Forest Products Studies 16. 559 pp.

 

1965. Genetic variation among 140 half—sib
Scotch pine families derived from 9 stands.
Silvae Genetica 12:75—106.

1965. Outline cooperative region project. 24 pp.
(Mimeographed).

., and W. 1. Bull.
1965. Geographic variation in Scotch pine,
results of a 5-year Michigan study. Silvae
Genetica 12:1-25.

., Walter L. Lemmien, and John Bright.

1965. Geographic variation in eastern white pine
——6—year results. Quart. Bul. Mich. Agr. Expt.
Sta. 45:691-697.

., S. S. Pauley, R. B. Polk, J. J. Jokela, and
R. A. Read.

1966. Performance of Scotch pine varieties in
the north central region. Silvae Genetica 15:
101—140.

 

 

VITA
JAVAID A. AHSAN
Candidate for the degree of

Master of Science

Major professor: J. W. Wright

Outline of studies:
Major: Botany
University of Panjab, B. Sc., 1958
University of Panjab, M. Sc., 1960
Major: Forestry
University of Peshawar,1% Sc. Honours,1962

Michigan State University, M. S., 1967

Biography:
Born February 2, 1940 in Jhelum, West Pakistan
EMperience:
Lecturer, Islamia College, Kasur, West Pakistan
August 1960-—December 1960
West Pakistan Forest Service

October 1962——continued
Organizations:
West Pakistan Class I Forest Officers Association

Society of American Foresters

' .... .1. 4.113345“ - -

".
t J'-

 

 

 

Thesis title:
FOREST TREE BREEDING RESEARCH IN THREE REGIONS OF THE
UNITED STATES AND ITS APPLICABILITY TO WEST PAKISTAN

CONDITIONS

 

 

 

 

 

 

 

 

 

 

 

 

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