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Apple Breeding

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DOI: 10.13140/RG.2.2.19929.03689

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 Class Lecture 2021

Apple Breeding

Dr Mahendra Kumar Verma,

ARS (1995), Ph.D., FIAHS, FISNS, FSHSD, FISHSD
Principal Scientist (Hort)
Division of Fruits and Horticultural Technology
Indian Agricultural Research Institute
New Delhi, India
E-mail: [email protected]
www.iari.res.in
 Content
1. Species and cultivars,
2. Cytogenetics,
3. Genetic resources,
4. Blossom biology,
5. Breeding objectives, ideotypes,
6. Breeding methods -introduction,
7. Selection, hybridization, mutation breeding, polyploidy
breeding,
8. Rootstock breeding,
9. Improvement of quality traits, resistance breeding for
biotic and abiotic stresses and
10. Biotechnological interventions, achievements and
11. Future thrust in
 Taxonomy
Family : Rosaceae
Subfamily : Maloideae (with pears)
Genus : Malus (40 sp.)
Section : Malus
Series : Malus
Species : domestica Basic chromosome No. – x=17
2n= 34, 51, 68
❖The cultivated apple has complex genome consisting of 34 chromosomes per
diploid genome.
❖The origin of the basic haploid number of n=17 in Malus and other Pomoideae
has been attributed to hybridization between two remote ancestral types,
Prunoideae=n=8 and Spiroideae=n=9.
❖Has gametophytic self-incompatability, and it is controlled by a single multiallelic
‘S’ locus.
❖When the pollen S-allele matches one of the pistils S-allele, fertilization is
inhibited.
 Nomenclature
In 1753, Linneaus in his ‘Species plantarum’ used the name
Pyrus malus for the common cultivated apple.

Miller in 1768 proposed Malus as generic name for the apples including three
species.
M. sylvestris
M. Pumila
M. coronaria

Poiret, 1804 given the name

M. communis

Borkhausen, 1803 proposed the name

M. domestica

M. x domestica Borkh. is the proper technical

name for the cultivated apples, owing to its
interspecific origin.
 Genome
• In 2010, an Italian-led consortium announced they had decoded the
complete genome of the apple (Golden Delicious variety).
• It had about 57,000 genes, the highest number of any plant genome
studied to date and more genes than the human genome (about 30,000).

Distributions of protein similarities for duplicated genes in

duplicated segments compared with grape (red), poplar (green)
and apple (blue).
 Origin
❖Domestic apple arose some 8-12million years ago in refuse
fruit forests in Central Asia, Primarily in the Tien Shan Region,
an area extending from the current region of Kazakhstan,
through the Xinjian Province in China, and all the way west to
Uzbekistan (Juniper et al., 1999).

❖Modern apple was developed from M. sieversii, which was

hybridized with from eastern origin species like M. prunifolia,
M. baccata, M. sieboldii, and from west with M. turkmenorum,
M. orientalis (Caucasian apple), M. sylvastris (Crab apple).

❖Molecular markers utilizing RAPD along with phylogenetic

tree analysis have confirmed that M. sieversii is the primary
progenitor of cultivated apple (M x domestica) (Lamboy et al.,
1996 ; Zhou and Li, 2000)

❖At one time each country and area had its own local cultivars.
 Diversity in Malus

Fruit Bloom
Fruits of M. sieversii

Late leaf fall

Bloom Dormant
M. baccata Early
“Kansas 14”
leaf fall
 Ancient Malus species of Mammal disseminated
M. Sieversii of Central
China: bird disseminated to Central Asia
Asia

The origin of the cultivated apple

North
America
became a
secondary
center of
origin: ‘Red
Delicious’,
‘Golden
Delicious’,
etc.

Johnny Appleseed
 1. Modern breeding objectives
Objectives for scion cultivars : Production of quality fruits for health and
prosperity

Major breeding objectives 1. To increase efficiency of the foliage.

reviewed by
Janick et al. (1996), 2. Structure of the tree to carry such
Laurens (1999), and weight
Brown and Maloney (2003)
3. Early bearing

4. Regular bearing

5. Good quality of the fruits

6. Good storage qualities

7. Disease and pest resistance

Objectives for rootstocks
Major breeding 1. Tree size control (dwarfism),
objectives
2. Tolerance to cold (hardiness),

3. Tolerance of pathogens and tolerance of

wet or dry soil conditions.

4. Long-term graft compatibility and good

health, e.g. freedom from viruses and

bacterial diseases caused by Agrobacterium

or Erwinia.
 Active breeding programs and important achievements of N. America
Country Organization Breeding objectives Important varieties developed /released/Introduced
USA Univ. of Arkansas - Stellar, Arkcharm
Univ. of Minnesota Cold hardy, high quality, Sweet Sixteen (1979), Honeycrisp (1991)
disease resistance
Cornell University, Fruit quality and Disease Cortland, Macoun, Empire, Jonagold, Liberty (1978),
Geneva, New York resistance Freedom (1983).

Purdue, Rutgers, Illinois Disease resistance GoldRush (1993),Enterprise (1993), Pristine (1995),
(PRI) cooperative Scarlrt O’Hara (2000).
Rutgers Univ., New Disease resistance Mollies Delicious (1966), Vista Bella (1974),
Jersey Suncrisp (1992).
Washington State Univ. Stress related to hot, dry and WA 2
sunny climate
Univ. of Florida Low chilling Anna, Dorsett Golden, Tropic Sweet
Canada British Columbia, Consumer preference and Spartan (1936), Summerred (1964), Scarlett Sentinel
Summerland sensory fruit quality (1997), Golden Sentinel (1997),
Quebec Fresh market, long storage, Belmac (1996), Primevere (1996), MacExcel (1996),
disease and insect resistance.
Nova Scotia, Kentville Scab resistance Nova Easygro (1971), Novamac (1978), Novaspy
(1986),
Manitoba, Morden Tree, shrubs, adaptability, Ornamental crab: Almey, Sundog, Garry, Selkirk,
landscape trade, Kelsey;
scion cultivars: September Ruby, Fall Red, and Red
Sparkle.
 Active breeding programs and important
achievements of Australia, NZ and Brazil
Country Organization Breeding objectives Important varieties developed
/released/Introduced
Australia Western Long storage Cripps Pink (Pink Lady) (1985), Cripps Red
Australia (Sundowner) (1985),
Queensland High quality red Apple thorpe Earlidel (1988), Apple thorpe
Horticultural apples, scab Summerdel (1989)
Institute resistance
Heverlee Transformation, Merlijn (King Jonagold x Liberty)
haploid induction,
molecular breeding
Brazil Empressa de Low chilling, Scab restant: Princesa (1986), Primicia
Paequisa disease resistance (1986);
Agro. Ext. low chill: Catarina (1996), Imperatiz (1997),
Rural Santa Baronesa (1997), Condessa (1997), Eva,
Catarina Anabela, Caricia.
New Zealand DSIR, Havelok Long shelf life and Gala, Splendour, Braeburn, Joy, Festival,
North fruit quality Pacific Rose, Southern Snap, Sci Early, Sci
Red
 Top Achievements

• Red Chief Red Delicious

Golden Delicious Oregon Spur

Red Chief Red Delicious Golden Delicious Oregon Spur

Gala Mast Mollies Delicious Gala Early Red One

Pink Lady CITH Lodh Apple-1 Chaubattia Princess

Ambri
 Scab Immune Varieties

Pristine

Redfree
 Active breeding programs and important
achievements of Asian countries
Country Organization Breeding objectives Important varieties developed /released/Introduced
China Hebei Changli Fruit Res. Institute High quality Yanshanhong (Ralls Janet x Richared Delicious (1989)
Zhengzhou Fruit Res. Institute High quality Huaguan (Golden Delicious x Fuji) (1988), Huashuai (Fuji x
Starkrimson) (1988)
Liaoning Fruit Research Institute All of the advantages Golden Delicious 463 (russet resistant)
on Fuji,
Shenyang Agricultural University Hanfu (Dongguang x Fuji)
Res. Inst. of Pomology (CAAS) Qiojin (1975)
Shaanxi Fruit Res. Inst., Yangling Quinguan (Golden Delicious x Jiguan) (1970)
Greece The Nat. Agric. Res. Found. Pom. Nauossa, Makedoni
Inst., Naoussa
SKUAST (K), J & K Fruit quality, shelf Lal Ambri, Sunehari, Akbar, Firdous, Shreen, Ambred, Ambrich,
India YSPUHF, H.P. life, high colour, early Ambstarking, Ambroyal, Chaubattia Anupam, Chaubattia Princess,
HETC, Uttarakhand ripening and scab Swarnima, CITH Lodh Apple 1, Pusa Gold, Pusa Amartara Pride
CITH, India resistance
IARI, RS, Shimla
Japan Aomori Apple Experiment Station Self compatibility
Self-compatible: Megumi (1950), Mutsu (1949), Sekaiichi
(1974), Hokuto (1983), Natsumidori (1983), Mellow (1990);
Self-thinning: Aori 9
Akita Fruit Tree Exp. Sta. Senshu (1983), Akita Gold (1990)
Gunma Agri. Res. Centre Akagi (1973), Yoko (1981), Sinsekai (1988), Gunma Meigetsu (1991).
NIFTS, Marioke Fruit quality, storage, Fuji (1962), Akane (1970), Iwakami (1985), Sansa (1986), Kizashi
disease resistance (1991)
Korea Nat. Hort. Res. Inst. RDA Storage life Hongro (1988), Kamhong (1992), Seokwang (1996)- early, attractive
(South) colour

Russia ARIH & ARRIGBFP, Michurinsk Skala

Chaubattia Princess CITH Lodh Apple-1
 Active breeding programs and important
achievements of European countries
Country Organization Breeding objectives Important varieties developed /released/Introduced
Czech Research & Breeding Institute Scab polygenic resistance Angold, Julia, Produkta, Zuzana, Jarka, Julia, Resista, Selena
Republic of Pomology, Holovouzy
England High quality, long storage, Fiesta (1984), Columnar (1991): Maypole, Tuscan, Trajan,
resistant to mildew, scab Charlotte, Obelisk.

France Institut National de la Resistance to pest, disease, Processing and Juice making: Jurella, Judaine, Judor;
Recherche Agronomique, reduction in orchard labour, scab resistant: Chanteline;
Angers improved fruit quality Others: Chantecler, Florina (1985), Baujade (1992),
Initial (triploid).

Delbard International Delbard Jubilee, Delearly, Harmonie Delorina, Delblush,

Nursery, Commentry Tentation, Regali Delkistar.

Germany Federal centre for Breeding Ahrina (1989), Gerlinde, Ahrista, Ahra.
and Research, Ahrensburg
Dresden-Pollnitz High quality, regular, Piros, Pilot, Pingo, Pirella, Pinova, Remo, Rewena, Rebella,
disease resistance Reanda
Greece The Nat. Agric. Res. Found. Nauossa, Makedoni
Pom. Inst., Naoussa
Italy University of Bologna Scab & mildew, resistance, Prime Red (Prima x Summerred) (1999)
compact habit, high fruit
quality
Experi., Insti., Fruit Culture, Quality, pyramiding of Red Earlilib, Golden Mira, Brina, Nova, Summerfree, Golden
Trento resistance Orange
The CPBO-DLO, Wageningen Quality, scab, mildew Elstar, Elan, Elise, Ecolette CPRO, Santana CPRO, Bellida
Netherlands resistance
 Prima Redfree Williams’ Pride

Sundance Pristine Enterprise

GoldRush Pixie Crunch CrimsonCrisp

 Prima Redfree Williams’ Pride

Sundance Pristine Enterprise

GoldRush Pixie Crunch CrimsonCrisp

 Early Introduction in India
Tamilnadu:- Perhaps the first ever exotic introduction of apple were made 1820 at
the Botanical Garden, Ooty (Tamilnadu) and later planted at Kodaikanal.

H. P.: First apple orchard was established at Bandrole in Kullu Valley by Capt. A. A. Lee
around 1860. Thereafter, many orchards were developed at Manali, Raison and Ngara by the
Englishmen.

A number of varieties were introduced by Alexander Coutts in his orchard at Mashobra in 1887.

Popular Delicious varieties were introduced by S. N. Stoke, a resident of Philadelphia, USA in

1918 at Kotgarh in Shimla Hills

Uttarakhand: Apples were introduced in Kumaon and Garhwal region by Britishers in the
later half of 19th Century after accession of this area in 1815 by defeating the Gorkha ruler of
Nepal.

Apples were introduced by Mr. Allen and Mr. Smith into Kumaon Hills at Chaubatia around 1872.
The first apple orchard was established in Ramgarh area of Nainital District.

Later many orchards were planted in Ramgarh, Bhowali and Hartola areas.
 Achievements from Controlled Hybridization in India
Hybrid Parents Breeding objective Institute
Lal Ambri Red Delicious x Ambri Yield and fruit quality SKUAST (K), Srinagar, J&K
Sunehari Ambri x Golden Delicious Yield and fruit quality SKUAST (K), Srinagar, J&K
Akbar Ambri x Cox’x Yield, scab resistance and fruit quality SKUAST (K), Srinagar, J&K
Orange Pippin
Firdous Golden Delicious Scab resistance, Yield and fruit quality SKUAST (K), Srinagar, J&K
x Rome Beauty x Prima
Shreen Lordlamborne x Melba x R- Yield, scab resistance and fruit quality SKUAST (K), Srinagar, J&K
12740-7A
Shalimar Apple-1 Sunhari X Prima resistance to scab, SKUAST (K), Srinagar, J&K
yields about 95 kg/tree (23.75 t ha-1)
Shalimar Apple-2 Red Delicious x Ambri tolerant to SKUAST (K), Srinagar, J&K
scab and Alternaria leaf spot,
Ambred Red Delicious x Ambri Yield and fruit quality YSPUH&F, Nauni (Mashobra),
Ambrich Richard x Ambri Yield and fruit quality Yield and fruit YSPUH&F, Nauni (Mash bra),
quality
Ambstarking Starkign Delicious x Ambri Yield and fruit quality YSPUH&F, Nauni (Mashobra),
Ambroyal Starking Delicious x Ambri Yield and fruit quality YSPUH&F, Nauni (Mashobra),
Chaubattia Early Sh. x Red D. Early maturity, Yield and fruit quality HETC, Ranikhet, Uttarakhand
Anupam
Chaubattia Early Sh. x Red D Early maturity, Yield and fruit quality HETC, Ranikhet, Uttarakhand
Princess
Swarnima Benoni x Red Delicious Early maturity, Yield and fruit quality HETC, Ranikhet, Uttarakhand
CITH Lodh Apple Bud sport of Red Delicious Yield and fruit quality with improved CITH Res. Stat., Mukteshwar,
1 overcolour Nainital, Uttarakhand
Pusa Gold Golden Del. X TEW Resist. Powdery mildew and apple scab IARI, RS, Shimla
Pusa Amartara Royal Del. X Prima Resist. Powdery mildew and apple scab IARI, RS, Shimla
 Methods of Improvement
✓ Introduction

✓ Selection (Clonal and seedling selection)

✓ Controlled hybridization

✓ Mutation and chimeras

✓ Marker assisted selections

✓ Transgene technology
• Majority of cultivated apple are diploid=2n=34.

• Darlington and Moffett (1930)- they are complex polyploids,

being partly tetraploid, partly hexaploid.

• Sax (1931)- apple is allopolyploid derived from a double hybrid

between two remote ancestors, one with the basic chromosome
of 8 and other 9.

• There are also triploid (2n=51) arises naturally from fertilization

of unreduced gametes= 10% of all apple cultivars. Important are
Baldwin, Gravenstein, Rhode Island Greening, Bleinheim Orange,
Mutsu.
 Breeding through Selections
Open-pollinated, chance seedlings proved to be outstanding cultivars,
including:
o‘Golden Delicious’,
o‘Delicious’,
o‘Jonathan’,
o‘McIntosh’,
o‘Rome Beauty’,
o‘York Imperial’,
o‘Stayman Winesap’,
o‘Yellow Newtown’,
o‘Winesap’,
o‘Rhode Island Greening’,
o‘Northern Spy’, and
o‘Gravenstein’.
oThese unique selections selected from thousands of growers became the
starting point for hybridization programs in the early history of apple
breeding.
 Development of Red Delicious and Golden Delicious
(Chance seedlings)
• The Red Delicious originated at an orchard in 1880 as "a round, blushed
fruit of surpassing sweetness.

• Stark Nurseries held a competition in 1892 to find an apple to replace

the Ben Davis apple. The winner was a red and yellow striped apple sent
by Jesse Hiatt, a farmer in Peru, Iowa, who called it "Hawkeye".

• Stark Nurseries bought the rights from Hiatt, renamed the variety "Stark
Delicious", and began propagating it.

• Golden Delicious, purchased from a farmer in Clay County, West

Virginia, in 1914.
 Breeding through Hybridization
• Controlled sexual hybridization among selected parents become
popular in 19th centaury, takes 5-7 years to complete a
generation.

• Breeding efforts made more progress when traits are known to

be controlled by qualitative rather than quantitative genes.

• Most traits, particularly those associated with fruiting and fruit

quality , are polygenically controlled.
 Inbreeding
• Apple have high degree of self-incompatibility.

• Difficult inbreeding

• Average fruit set from selfing is about 2% & no. of seeds/fruits is 1-2.

• Only 30% of seeds are able to germinate.

• Seedlings are weak and may die.

• Minimum 3-generations are required to produce few homozygous lines,

which increases further in next generation
 Out breeding and Backcrossing
• Most commonly done.
• This is important when we require to introduce one character of importance
e.g., disease resistance.
• Therefore, modified backcross method is recommended by Hough et al., 1953
to overcome the demerits of inbreeding.
Example:- Wild Spp. X Large Fruited cultivar
F1 (Heterozygous)= only best are selected

F1 x Cultivar

(produces 50% resistant plants)

Best are x good cultivar

(until all are resistant)
 Development of varieties through hybridization

Flowers are collected at Pollen collection:-Remove the

the balloon stage just before
the petals expand, and before anthers by rubbing with small comb, collect in
the flower dehisce.
Petri dish, keep in sunny side

Harvestin
g of fruits
Store in
refrigerator for using
and Stratification
within one month or
removal of
store at -15 in deep
seeds
freeze for one year

Cross Emasculation of Viability test Quality testing

Pollination: female parents, remove on a drop of with
dip brush in petals, sepals, stamens sugar acetocarmine –
vial and with nails of thumb and solution (2.5- glycerol (Marks,
touch the index finger before buds 20%), or 10% 1954) ,
same on to reaches balloon stage- 2 sucrose at determine under
stigma flowers per cluster 20-250 C microscope
 Stratification of seeds Juvenility:- period between germination to start of
fruiting
3-50C ; 6-14 weeks
Leaves are smaller
Finely serrated
Shoots are thinner
Produced at right or obtuse angle
No flowers produced

Transition phase Onset of adult phase

Development of flower buds

Lower part is still

juvenile
& upper part is adult
 Floral Biology

❖Flower colour- White to ❖ Pollen produced in great

deep pink excess
❖Produced in the mixed bud
❖ 20 anthers
in cymose clusters on spurs
❖Sepals-5 ❖ 3,500 pollen grains /
❖Petals-5 anther
❖Stamen-20 ❖ 70,000 pollen grains /
❖Pistil-1 divided in to 5 style flower
❖1-Ovary, has 5 carpel's
each containing 2-ovules ( ❖ 5 stigmas / flower
Maximum seed number is each with 2 ovules
10) ❖ Approx. 7,000 pollen
❖Few cultivars have more grains / ovule
than 10 up to maximum of
20.
 • Apples require cross pollination
•
Pollination •
Careful selection of pollen provider trees is necessary
– Pollen providers called “pollenizers”
Not all apple cultivars produce pollen that will fertilize all
other apple cultivars
– Example: Winesap pollen is sterile
to
other
flowers 2.Pollination (pollen transfer to a stigma)

Pollination:-
1.Anthesis (pollen shed) Mechanical
transfer of
pollen from
3.Pollen germination (on stigma) anthers to
stigma.
Carried out by
insects-bees,

4.Growth of germ tube to ovary

For making hybrids

5.Fertilization of ovule controlled pollination is
required
 Ways of shortening juvenile period
• Ways of shortening the duration of the juvenile phase have been investigated
(Kemmer 1953; Murawski 1955; Visser 1964).

✓ Methods to induce adult trees to flower include shoot priming, root priming, and
bark ringing.

✓ Grow the seedling as fast as possible in glass. Seedlings flower in the greenhouse
in 1- year by keeping them in a continuous state of growth (Aldwinckle et al.
1976) under optimum conditions in the greenhouse, closely planted, watered,
and fed, may reach 3 m in height in the first season.

✓ Way (1971) was able to induce seedlings to flower by bark ringing when they
were 4 years old, but 3-year-old seedlings did not respond.

✓ Tydeman and Alston (1965) shortened by budding seedlings onto dwarfing

rootstocks.

✓ Juvenility poses a problem for the rootstock breeder. In the stoolbed, juvenile
plants are usually quite thorny, root very easily, and exhibit earlier bud break and
later leaf fall.
 Early selection
• Specialy disease infection through leaves such
as scab, powdery mildew
• The plant growth parameters
 Preselection criteria
Any positive correlation of this kind which is helpful to discard in
the nursery stage example:-
1. pH of leaf sap the sweet type usually have pH 5.7 and the
acid type 5.5-5.6- not significant.
2. Colour of the fruits and anthocynin pigmentation of one year
shoot and petiole.
3. Late leafing and late flowering.
 Evaluation for the fruits

✓Fruits > 65 mm (70 mm as per codex standard) diameter

✓Good colour e.g., clear yellow (Golden Delicious), bright red

(Red Delicious, Oregon Spur, Red Chief, Red Fuji), bright
green (Granny Smith)

✓Good apple flavoured (Golden Delicious, Ambri)

✓Sub-acid with pleasant flavour (Mollies Delicious, Gala Mast

etc)

✓Firm texture, plenty of juice (Fuji and Gala and their strains)
 Records

• Its most important part of any breeding

program.
• Parameters- tree growth, flowers, fruit
characters etc.
• IBPGR Passport information sheet as well as
DUS guidelines developed by PPV & FRA as
well as UPOV.
 Characters controlled by single genes
• Very useful
• Anthocynin pigmentation discovered first in M. pumila.
• Easy to transfer if controlled by a dominant gene.
• 5-6 generations are required to eliminate the
undesirable characters and retain the good qualities of
large fruited cultivars.
• If the character is controlled by recessive gene-
required more generations.
 Characters controlled by polygenes
• Most of the quality traits are governed by polygenes
e.g., fruit size, shape, cropping etc.
• Progeny means and parental means are important
which depends on degree of dominance of the
particular character
 Selection of Parents
• Based on the object of the breeding programme
• Either both parents must have the dominant character or
one must be.
Example:-
Resistant to Scab : Antinowka, M. baccata, M. sargenti, M.
seiboldii, M. zumi calocarpa, M. floribunda
Podery mildew (Podosphaera leucotricha)resistant:-M. robusta,
M. zumi.
Collar rot resistant (Phytopthora cactorum):- MM.103
Wolly aphis (Eriosoma lanigerum) resistant:-Northern Spy
 Mutations and chimeras Breeding
• A bud sport or mutation is a variant of an
inherited character arising in a cell from which a If the mutation
bud eventually develop; this in due course is limited to
produces a shoot and later a branch which only one cell
differs, usually in one character, from the plant layer are
on which it was produced. chimera; since
the gametes
• Two types of mutations in apple:-
are formed
• 1. Those which differs only single gene of tree from the sub-
or fruits. epidermal
• 2. Those which alter the ploidy layer
• Varieties Most prune to bud sports:- Delicious,
Rome Beauty, Winesap, Cox’s Orange Pippin
 Triploids
• More vigorous
• Larger fruits
• Produces very poor fruits- leads to pollination
problems-requires a diploid to pollinate the
triploid.
• Produces very few good seeds
• Useless as parent for further breeding-
produces if selfed or crossed very few seeds-
are mostly aneuploids seedlings.
 Tetraploids (2n=68)
• Discovered as large fruited sports on trees of diploid
cultivars.
• Usually occur as chimeras with one or more layers of
diploid tissues over tetraploid.
• If one diploid layer occur, tree behaves as a tetraploid in
the formation of gametes; with more than one diploid
layer, it behaves as a diploid.
• Fruits are misshapen, and large.
• growth short-jointed,
• Cross between tetraploid and diploid give large progenies
of triploids.
 Marker-assisted Selection
• Efforts are underway to map the genomes of many fruit species with special
emphasis on the Rosaceae (Folta and Gardiner 2009).

• Full genomic sequencing in peach, apple, and strawberry has made it

possible to locate useful genes include those affecting quantitative traits.

• The combination of advances in informatics, sequencing technology, and

genomics makes it theoretically possible to select on the basis of the
genotype instead of the phenotype.

• This has potential benefits to fruit breeding where the cost of phenotype
selection are very high because of the long time necessary for fruiting and
the large amounts of land required for each seedling.
 Plant Genetic Engineering
TRADITIONAL PLANT BREEDING
DNA is a strand of genes, much like a strand of pearls. Traditional plant breeding
combines may genes at once
Many genes are
transferred
Desired Gene

X
Commercial New
Traditional Plant Variety Variety
Donor

PLANT BIOTECHNOLOGY
Using plant biotechnology a single gene may be added to the strand
A single gene is
transferred
Desired Desired
Gene Gene

+ Improved
Commercial Commercial Plant
Donor Plant Variety Variety
 Schematic Illustration of Steps involved in
Generating Transgenic Plants in the Lab

Explants Heterogeneous
embryogenic
Transgenic Plant calli
Agrobacterium
transformation Particle
bombardment

Southern Western Enzymes

Molecular analysis and verification
Selection on
suitable medium

Plant regeneration Putative transformed Selection of

(with/without selection) embryogenic calli transformed calli
 Development of Transgenics
◆ Identify gene(s)
giving a desired trait
◆ Make copies of the gene
◆ Transfer to plant tissue
◆ Regenerate plants
◆ Lab analysis and safety testing 7-8 years
◆ Development of a variety
◆ Field tests
◆ Approval by Government agencies
◆ Commercialization
Salient Steps involved in the Development of
Transgenic Plants from Lab to the Field
Transgenic Plants Confirmed in Lab

Green house & Limited Field Tests

Ecological & Biosafety Analysis

Regulatory Permit for Large-scale Use

Proprietary Arrangements

Commercial Deployment

Continued Monitoring of Efficacy and Safety

 The development of a cisgenic apple plant.
Vanblaere et al. (2011), J Biotechnol. 2011;154(4):304-11.

• Cisgenesis is toward a new generation of GM crops.

• Insert of gene(s) derive from organisms sexually compatible crop.
• Exa. Insertion of the endogenous apple scab resistance gene HcrVf2 under the
control of its own regulatory sequences into the scab susceptible apple
cultivar Gala.
• Three cisgenic lines were chosen for detailed investigation and were shown to
carry a single T-DNA insertion and express the target gene HcrVf2. This is the
first report of the generation of a true cisgenic plant.
 Breeding specific characters
1. Breeding for vigour
• Dwarfing stature is desirable for rootstocks.
• Weak progenies can be produced by inbreeding &
• Aneuploids
Triploids X Triploids
Triploids x Diploids
Triploids x Tetraploids
 2. Cold hardiness
• Useful to extend cultivation in colder areas.
• To protect from winter injury.
• The breeding and selection for cold hardiness in deciduous fruit
crops is reviewed by Stushnoff (1972).
• Portable low temperature chambers are used for study.
• Stuart (1941) & Wilner (1960) gave a method to measure the
electrolytes exuded by leaves when subjected to undergo cold
treatment. These electrolytes are measured by electrical
conductivity meter.
• Lapins (1962) also given a method- one year old shoots are
exposed to cold from 00 to -430c lowered by 20c/ha. Recovery of
the shoots observed after a week (1-10 point scale)
 3. Spur Type
• Compact habit, reduced internodal length, limited side
branching, prolific development of fruit spurs,
precocious, heavy crop, regular bearing- dwarfs or
compact mutants.
• This is controlled by single recessive gene.
• X-ray irradiation of dormant scion (3Kr)- grafting---------
----shoots is cut back in to original grafts to induce
latent buds----------process is repeated for 3 years.
• By controlled crossing between Mutants of McIntosh X
Golden Delicious produces 43.9% spur type progeny.
4. Fruit size
Most critical character.
Accepted size is 65mm dia. And large size is 75mm
Controlled by polygenes.
5. Fruit shape
Very flate to oblate.
Polygenic control.
6. Seasons of ripening
August is Early; March is Very late (0-7)
 7. Fruit colour
• Determined by the ground colour of the skin and
anthocynin pigmentation of the skin.
• Ground colour the polygenically controlled.
• Anthocynin is controlled by dominant gene.
• Anthocynin present or absent; if absent, either fruit
colour is yellow or green.
• If pigmentation is present – from small red flecks to bold
stripes and from faint blush to solid red.
• Anthocynin pigment is in solution in cells of epidermis and
two sub-epidermal layers but not all cells are pigmented..
• Intensity of colour depends on the proportion of cells in
each layer.
8. Fruit Flesh colour
• White-cream-pale yellow or
• Green white- tinged with red.
• Preferred colours are clear yellow and red skin colour
with white or creamy white flesh.
9. Russet
• Varies from completely russet to free of russet.
• Genetical and environmental behaviour.
• Environment like low temp., humidity, spray damages
common in Golden Delicious which is susceptible to
environment.
 10. Apple scab
• Two form- polygenic and monogenic
• Most desirable types of resistant can be obtained with
a combination of both.
• Antonowka- have field resistance to all known races of
fungus are of polygenic types.
• M. floribunda- has single dominant gene (Vf) resistance
against scab.
• Backcross method is used which produces 50% resistant
progeny in each generation (Prima and Priscilla
developed by Dayton et al., 1970)
11. Mildew
• Caused by Podosphaera leucotrica.
• Inheritance is quantitative, but without the
knowledge of dominance.
• Single gene (Pl1) dominant immunity found in M.
robusta & M. zumi (Pl2)
12. Collar Rot
• Caused by Phytopthora cactorum.
• Controlled by polygenic and monogenic
inheritance.
13. Woolly Aphis
• Caused by Eriosoma lanigerum
• Controlled by quantitative genes
• Northern Spy has single dominant gene (Er)
 14. Rootstocks
• Takes longer time to assess
• Good rootstock must have clean upright stem for easy
to bud or graft,
• good roots to provide better encourage,
• stock-scion compatibility,
• sucker free,
• precocious,
• heavy crop support
• Resistant to crown gall, woolly aphis, canker, mildew etc
• And high degree of winter hardiness.
 Problems in
apple breeding
 Sterility and Incompatibility
• Two main causes –Sterility & incompatibility.
Sterility:-
• Due to failure in the any of the process of pollen,
embryo-sac, embryo and endosperm development.
• Is due to chromosomal imbalances e.g., Triploidy-
leads to very poor seed set and poor pollen
production.
• Exception:- in diploids pollen abortion occurs due to
lethal genes.
 Mechanisms to prevent self-pollination:
Sexual incompatibility
– compatibility genes must have different alleles for pollen
to germinate

Gametophytic compatibility
 Incompatility
Sexual Incompatibility
❑Due to failure of the pollen (although are
functional) to grow down the style & do
fertilization- common in apple.
❑Therefore, all the apple cultivars are practically
self-incompatible.
❑Gametophytic incompatibility.
❑Selfing causes 0-9.6% fruit set only e.g., selfing
in Cox’s Orange Pippin, Golden Russet &
Northern Spy gave only 1.5% fruit set.
 Apomixis
Production of seeds from unfertilized eggs, which never
produces true seeds.
• All are polyploids Exa:- M. sikkimensis, M. coronaria,
M. toringoides, M. sargentia, M. seiboldii.
• In normal course these produces apomictic seeds, but
if crossed with diploid- produces sexual hybrids.
• Uses:- Seedlings produces are uniform, virus free and
can be used as rootstocks.
• Inheritance:- this has dominant trait.
 Parthenocarpy
• Development of fruits without fertilization (Parthenogenesis) ;
whereas, Production of seeds from unfertilized eggs, which
never produces true seeds(Apomixis).
• Normally fruits have ten ovules- but not necessary that all seeds
are necessary to develop the fruits; a single seed is sufficient to
produce the seed in apple.
Artificial induction of parthenocarpy:-Can be induced by
Gibberellins A3 and A4 (5 x 10-3 and 5 x 10-2 M) induced
parthenocarpy .
Characters of Parthenocarpic fruits are:- are good where inclement
weather conditions occur like late frost.
I. Fruits are small & misshapen
II. Ripens early
III. Short shelf-life
 Clonal Rootstocks
✓It was early as 3oo years before Christ that the Alexander The Great sent dwarf
apple trees back to Greece from his conquests of Asia.
✓The ‘ Paradise’ which was named to a group of dwarfing rootstocks of apple
perhaps selected from Persia, Paradise and Golden of Eden.
✓Thereafter, due to large number of clonal rootstocks were grouped in to two
categories
✓ Paradise stocks:- hairy rooted having severe dwarfing effect.
✓ Doucin Stocks:- semi-dwarfing rootstocks.

✓ Example:- Holly Leaf, Holstein, Doucin, English Broad Leaf, June-de-Mtz etc.

The clonal rootstocks, propagated asexually, provide options in meeting the

demands of modern orcharding such as uniformity, size control, precocity, cropping
efficiency and resistance against certain adverse biotic and abiotic factors. The
cultivation of apple has been shifted almost entirely to one or other of these clonal
rootstocks all over Europe, USA, Canada and other apple-growing countries of the world.
Malling and Malling Merton rootstocks

• First successful effort to test and evaluate rootstocks was initiated at East Mailing
Research Station, Kent, England, in 1912, which resulted in the development of
Malling series of clonal rootstocks. The first series consisting of 16 stocks was given
Roman numerals as “Malling I” and later on abbreviated to M I, M II and so on.
Subsequently been raised to 27.

• First 25 were numbers in Roman numerals and all others beyond 25 are given
Arabic numerals M 26.
• They were: M25 in 1956, M26 in 1965 and M27 in 1976, raising the total number to
27.
• During period of their popularization, Australia and New Zealand facing the serious
problem of woolly aphid on the roots of apple trees. Mallling series were not
found of any help.
• East Malling (HM Tydeman) and John Inns Research Institute (Merton) in England
a total of 15 rootstocks were numbered from MM 105 to MM115 in 1928.
 S. No. Resultant Parents
Progeny
1 MM. 101 Northern Spy x Malling I
2 MM. 102 Northern Spy x Malling I
3 MM. 103 Northern Spy x Bendavis
Original botanical
4 MM. 104 Malling II Northern Spy
description of these
rootstocks have been 5 MM. 105 Malling II X Northern Spy
described by Tydeman 6 MM. 106 Northern Spy x Malling I
(1953) and behaviour 7 MM. 107 Northern Spy x Malling XV
summarized by Perston 8 MM. 108 Northern Spy x Malling XV
(1953). Gammer(1953)
9 MM. 109 Malling II X Northern Spy
reported propagation
qualities of these crops 10 MM. 110 Northern Spy x Malling I
11 MM. 111 Northern Spy x Merton 793 / Northern
Spy x Malling II
12 MM. 112 Northern Spy x Winter Majestin
13 MM. 113 Northern Spy x Malling XII
14 MM. 114 Northern Spy x Malling XII
15 MM. 115 Northern Spy x Bendavis
• During the recent years, EMLA series of rootstocks, EM from East Malling and LA
from Long Ashton, have been introduced. The EMLA rootstocks, which are
designated as EMLA9 or M9 EMLA and EMLA106 or MM106.
• EMLA, are nothing but M or MM series of rootstocks which are free from known
virus diseases prevalent in apple plantations in the United Kingdom. The new
EMLA9, however, is much more vigorous than M9 and appears to be a different
strain or clone, rather than simply virus free M9.
• In India, elaborate rootstocks trials were initiated on M and
MM series of apple rootstocks soon after their introduction
during sixties at Mashobra and Kotkhai in Himachal Pradesh,
at Shalimar in Jammu and Kashmir and Chaubattia in
Uttarakhand.
• These rootstock trials generated useful information on the
performance of different clonal rootstocks under varying
agro-climatic conditions.
• Many promising rootstocks have been identified and
recommended for commercial use in different states. These
included M7, M9, M26, MM106, MM111 in Himachal
Pradesh, M2, M4, M7 and M9 in Jammu and Kashmir and
M13 and MM106 in Uttarakhand.
 Ideal Rootstock
• Freestanding • Disease & Insect
resistant
• Precocious
• Widely adaptable
• Dwarfing
• High Yielding
• Graft compatible
• High quality
M2
• This is a vigorous rootstock.
• Producing a tree slightly larger than M7 but smaller than seedling.
• Precocious and productive.
• Adapted to most soil types but is not suitable for heavy soils.
• It is resistant to collarrot, but susceptible to root-rot.
• This rootstock does not propagate as easily as MM106.

M4

• It produces a semi-vigorous tree slightly larger than a tree on M7 and slightly

smaller than M2.
• It induces earlier and heavier production than M2, and heavier and more regular
production than M7.
• This is not well-anchored as the roots develop mainly on one side. The plants on
this rootstock need staking.
• Though susceptible to water logging, it is comparatively resistant to collar-rot.
• It is easy to propagate through mound layering.
M7
• A semi-dwarfing rootstock, it produces a tree 55-65 per cent the size on
apple seedlings.
• It is adaptable to a wide range of soil types and climates.
• It can withstand temporary drought as well as waterlogging.
• Suckering is a problem in this rootstock, but high budding and deep
planting reduce suckering
• This is susceptible to root-rot, but less susceptible to collar-rot.
• Since this propagates easily, it is recommended for nurserymen.
M9
• As dwarfing rootstock it is most widely used throughout the world.
• It is most suitable for high-density planting.
• That is why it is used most extensively in Europe.
• Its roots are shallow and brittle, and hence the anchorage is poor and
trees on M9 need support.
• It cannot stand water logging and drought.
• it is resistant to collar-rot.
• M9 is very difficult to propagate through hardwood cuttings, though it
roots fairly well in stool beds.
M 13
• A vigorous rootstock, it produces a full size tree similar to apple seedlings.
• It is more tolerant to wet soil conditions than other rootstocks.
• M 13 roots easily, but it produces a spreading shallow root system with cultivars.
• Therefore, it is useful in shallow soils or soils underlain with hardpan.

M 26
• A cross between M9 and M16.
• it produces a dwarf tree slightly larger than M9.
• It is more precocious and productive than M9.
• It is useful for high-density planting in soils where M9 performs poorly.
• It is non-suckering but the trunk forms bur-knots. Therefore, it should be planted
with graft union as close to the ground as possible.
• M26 is susceptible to fire blight and collar-rot, but is most winter hardy of the
Mailing rootstocks now used commercially.
• This rootstock is easily propagated by hardwood cuttings but is a poor producer
in stool beds.
MM106
• A semi-dwarf rootstock, it produces a tree slightly larger than M7.
• Trees on this rootstock are very productive, well-anchored and do not sucker.
• Fruit bearing starts early on MM106.
• It is resistant to Woolly Aphids and seedling blight, but susceptible to powdery
mildew and collar rot.
• It is easily propagated through softwood and hardwood cuttings as well as
through stool layering.

MM111
• A vigorous rootstock, it produces a tree about the same size as that of M2.
• It is not precocious but it is much more productive than apple seedlings.
• It is adaptable to a wide range of soils and climatic conditions.
• It is highly tolerant to drought and resistant to collar rot but susceptible to
powdery mildew.
• It is easily propagated by stooling.
Other clonal rootstocks

Polish apple rootstocks

• The Polish apple rootstocks (P series) have been developed with the principal
objectives of vigour control and tolerance to winter cold.
• Among the first series (P1, P2, P16, P18 and P22) have been found promising. Of
these, P22, P2 and P16 are the most important.
• These 3 clones have been raised from crosses of M9 with Antonovka. All are
winter hardy and resistant to collar (crown) rot. These induce good yield
precocity and efficiency in the scion. However, these are susceptible to fire blight
and Wooly Aphids.
• The P22 and P2 are as promising as dwarfing rootstocks and interstocks. Trees on
P22 are extremely dwarf, while those on P2 are slightly more vigorous than those
on P22.
• A second series of Polish apple rootstocks, P59, P60 and P92, have been released
more recently, but these have not been tested extensively so far.
Russian apple rootstocks
• Two types of apple rootstocks have been developed in Russia. Budagovsky (Bud
or B) series from and the Krasnodar clones (KC).
• The Budagovsky (Bud or B) series of rootstocks are cold hardy, providing
adaptability to severe climate of Central Russia and cover full range of size
control. Of the series, B9, B146, B490 and B491 are most important. B9, B146 and
B491 are dwarfing, while B490 is a semi-dwarfing rootstock. B146 is the most
dwarfing. Being similar or slightly more dwarfing than M27. B491 is between
M27 and M9 in vigour; while B9 controls the tree size to a level between M9 and
M26. B9 is the most extensively tested rootstock and its principal benefits are its
extreme cold hardiness and resistance to collar-rot.

• Among Krasnodar clonal rootstocks, KC-1 (formerly 1-48-46) and 1-48-41 clones
are important. These rootstocks were developed from crosses of M9 and M4
rootstocks with local varieties. Both these rootstocks are drought resistant and
semi-dwarfing. KC-1 increases yield over the Malling rootstocks.
American apple rootstocks
• The American apple rootstocks constitute a varied group of rootstocks that were
introduced from the breeding and selection programmes from Michigan, New
York, Missouri, Kansas and Oregon.
• The Michigan apple clone (MAC) series of apple rootstocks were originated from
open-pollinated seedlings from planting of Mailing rootstocks, Alnarp 2 and
Robusta 5.
• In this series, MAC1, MAC9 (Mark), MAC24 and MAC39 are more promising.
• MAC1 is semi-dwarf. The trees on this rootstock are similar in size to those on
M7, while MAC24 is semi-vigorous in vigour class of MM111. Mark (formerly
MAC9) and MAC39 are dwarf rootstocks.
• Mark is most popular, producing trees slightly larger to those on M9, but similar
to M26. Trees on Mark rootstock show excellent precocity, yield efficiency and
good anchorage.
• Mark is also easier in propagation than M9.
Geneva (G) series formerly Cornell Geneva (CG) series
• Developed at Geneva, New York, from McIntosh, Northern Spy and Malling clone
crosses.
• G11, G13, G16, G30 and G65 are promising.
• G11 is similar in size control to M26.
• G16 is same as M9 in size control, but it is more precocious than M9.
• G65 is half-way between M9 and M27 in size, but is precocious and productive. It has
good anchorage.
• G13 is little more dwarfing than M7, while G30 is similar in size to M7, but is
precocious and has excellent anchorage.
• These rootstocks are resistant to fire blight, G65 being almost immune to fire blight.
• G65 and G30 are also resistant to collar-rot.

Besides G rootstocks, Geneva New York Station also introduced one clone, Novole (P1
286613) in 1982, as a rootstock. It is also resistant to fire blight, crown rot and tomato
ring spot virus. It is easy to propagate by cuttings. Since it is a vigorous rootstock, it is
recommended for use with dwarfing interstocks.
C series of apple rootstocks
Developed by Stark Bros Nurseries in Missouri from the population of open
pollinated seedlings M8.

• Three selections, C6, C46 and C59 are promising.

• C6 was introduced in 1974 for use as dwarfing interstem tolerant of
common latent viruses. Tree size control is similar to that of M9, when
used as interstock or rootstock, and is similar in susceptibility to fire
blight and Wooly Aphids, but is less brittle than M9. It is compatible with
a wide range of cultivars, inducing very early and heavy production.

• K14 clone was selected at Kansas Agricultural Experiment Station in 1938

from a population of French crab that survived 2 intense summer
droughts and a severe freeze. Since it is a very hardy rootstock, it is used
as a hardy trunk stock or interstock.

• A dwarf clone, Oregon Apple Rootstock-1 (OAR-1), was selected for its
growth control and outstanding anchorage.
 Canadian apple rootstocks

• Ottawa series, Kentville stock clones and Vineland rootstocks.

• 2 types, Ottawa Hybrid seedlings (OH) and Ottawa clonal (0) series.

• In OH series, 6 rootstocks (OH-1 to OH-6) were introduced in 1971. All these

rootstocks are vigorous and tolerant to latent viruses commonly found in
commercial apple cultivars.

• In Ottawa clonal (0) series, originally 10 clones (1 to 10) were selected of which
Ottawa 3 and Ottawa 8 are promising. Ottawa 3 is between M9 and M26 in
vigour. The principal benefits of Ottawa 3 are its better anchorage and much
better hardiness than M9, but major constraint with it is extreme difficulty in
propagation. The Ottawa 8 is a semi-dwarf rootstock, producing trees equivalent
in vigour and productivity to those on MM106. It is the much hardier rootstock
than MM106.
Robusta 5 is an important winter hardy rootstock. Originated by the
Canadian Department of Agriculture in 1928 from the seed obtained
from Siberia, it is a vigorous rootstock. It produces trees equal in size to
apple seedlings. It is resistant to fire blight and easy to propagate.
 Swedish apple rootstocks
Alnarp2 (A2) and Bemali are important Swedish rootstocks. An
introduction from Alnarp Fruit Tree Station, Sweden.
Alnarp 2 is a winter hardy and vigorous rootstock. It induces early
bearing and productivity.
Bemali is an introduction from the apple breeding programme at
Balsgard, Sweden, from a cross between Monk's Golden and M4. It is a
dwarf rootstock with tree size in the range of M9 to M26. It is precocious
and productive. Its anchorage is better than M9, but not as good as M26.
Resistant to fire blight and Wooly Aphid, it is very easy to propagate.
 Other new rootstocks
• Apple rootstock breeding programmes are underway in several other countries France,
Germany, Belgium, Netherlands and Czechoslovakia.

• The principal objectives of these breeding programmes are: to develop apple rootstocks
that are free from shortcomings of the important current commercial rootstocks, and also
rootstocks which may be more vigorous than M27 and less vigorous than M26 and more
suitable for high density planting.

• The Jork Research Station, West Germany, has released Jork 9 (J9) rootstock. It is a
selection from population of open pollinated seedlings of M9. It is slightly more dwarfing
than M26, inducing equal or slightly more production than M9 and M26. It is easily
propagated in stool beds.

• The J.T.E. series of apple rootstocks from Czechoslovakia are under testing in many
countries. Of which, three clones (F,G and H) are promising. Clone G has vigour similar to
M27, clone F similar to M9 and clone H approximately 25 per cent greater than M9.

• French apple rootstocks, Pajam 1 (Lancep) and Pajam 2 (Capiland) are dwarfing and easier
to propagate than M9. Trees on Pajam 1 are slightly smaller than M9 EMLA and Pajam 2,
while Pajam 2 is similar to M9 EMLA in vigour.
5. Inter-specific hybrids
Colt Hybrid of P. avium x P. pseudocerasus developed at East Malling
Semi- dwarfing
Becoming popular commercially for high density plantation.
MxM (Mazzard x Mahaleb )Clones
These rootstocks are more precocious & productive than F 12/1
Tolerant to nematodes
M x M 14 Dwarfing rootstock
M x M2, M 39, M 60, M 97----- Resistant to Root rot
M x M 14, M 39,M 60,M 97---- Tolerant to Canker
 A) Drought Tolerant :
 Apple : MM 111, KC-1, KC1-48-41

B) Cold Hardy
 Apple : B 490, B 491, Bud-9(RLP), O-3, P2, P22, Novole,Alnarp 2.
C) High Temperature Tolerant
➔ Apple : M-7, MM-109.
D) High Soil pH Tolerant
➔ Apple : M-9, MM106,
F) High Soil Moisture Tolerant
 Apple: M7,MM104, MM106,.

G) Disease Resistant Rootstocks

i. Resistant to Powdery Mildew
 Apple : P1,P2,P16,P18,P22.
ii. Resistant to Latent Viruses
 Apple : B 9, C 6, MAC 9, Novole.
iii. Resistant to Crown & Root Rot
 Apple : B 9, B 491, G 30, G 65, MAC 9, O 3, P 2, Novole.
 The IARI Regional Horticultural Station, Shimla

• Collected a number of wild and indigenous species for their evaluation as rootstocks
for commercial cultivars.
• Of the 6 M. baccata types, M. baccata Rohru was observed to be more dwarfing than
M9.
• M. baccata Shillong, M. baccata Khrot, M. baccata Gaibong and M. baccata Dhak,
were as dwarfing as M9.
• The M. baccata Srinagar was observed to be semi-dwarfing, similar to MM106
rootstock.
• All these 6 types gave good grafting success with Golden Delicious apple. These were
easy to propagate by stooling.
• The Malus baccata var. Himalaica is as dwarfing as M9 and has good compatibility
with apple. It is resistant to collar rot, root rot and Woolly Aphid.
• The M. baccata var. Himalaica is used as a rootstock for apple in Uttaranchal and
appears to hold promise as a suitable rootstock.
 Breed it…………its beautiful

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