10/18/2019

Citrus Rootstocks

Ute Albrecht [email protected]

Southwest Florida Research and Education Center UF/IFAS, Immokalee, FL

PLP 5115C – Pathology Fall 2019

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Outline

I. Grafting II. Rootstock propagation III. Importance of rootstocks IV. History of rootstocks V. Rootstocks and HLB

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Definitions

. “Stionic” interactions = interactions between the stock and the scion. Scion . “Stion” = any or tree composed of a Graft union stock and a scion growing in combination. (Webber, 1932) Rootstock

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I. Grafting

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Grafting The process by which part of one plant (the scion) is inserted into another (the rootstock or stock) so that they unite and form a single plant. Budding A type of grafting with the scion consisting of a single bud.

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Citrus nursery program . Propagation material is obtained from the Division of Plant Industry (DPI), Bureau of Citrus Budwood Registration. . The Citrus Budwood Registration Program was started in 1953 to prevent diseases and ensure a healthy commercial citrus industry. . New material is cleaned up and tested if true-to- type before being released to growers.

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https://www.freshfromflorida.com/Divisions-Offices/Plant-Industry/Agriculture- Industry/Citrus-Health-Response-Program/Citrus-Budwood-Program

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Budwood trees

M. Zekri

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Budwood

(Webber 1946)

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Inverted T budding method

(Reuter 1973)

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Inverted T budding method

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Inverted T budding method

Inserting and wrapping the bud

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Bending of rootstock liners to push bud growth

M. Zekri

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Bud growth Finished trees

M. Zekri

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Graft union

Source: Webber (1948) The smoothness of the bud union was (is) thought to be indicative of the compatibility between rootstock and scion.

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Graft union

A B C

D EF G

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II. ROOTSTOCK PROPAGATION

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Rootstock propagation is traditionally by seed.

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Nucellar embryony Nucellar embryos. Embryos derived from nucellar tissue. Apomixis. Development of an embryo without fusion of female and male gametes. Polyembryony. Formation of more than one embryo in a seed.

. Vegetative (≠ zygotic) embryos. . Clones of the mother tree. . Start to develop earlier than zygotic embryos. . Different rootstocks have different degrees of nucellar embryony. YourArticleLibrary.com

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Implications of nucellar embryony

. True-to-type propagation of rootstocks. . Elimination of viruses. . Difficulty of breeding citrus as only few seedlings will be genetically variable.

(Koltunow et al 1993)

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Rootstock fruits and seeds Sour Carrizo

Swingle Cleopatra mandarin

http://www.citrusvariety.ucr.edu/citrus

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Alternative propagation methods

. Clonal propagation is also possible through use of cuttings or tissue culture (vegetative propagation).

. Both methods will yield genetically uniform .

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Alternative propagation methods

. Clonal propagation is also possible through use of cuttings or tissue culture (vegetative propagation).

. Both methods will yield genetically uniform plants.

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Seed source trees

Exposure to disease

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Seed source trees

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Seed source trees Diseases: HLB, citrus scab, , etc.

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III. IMPORTANCE OF ROOTSTOCKS

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Why use rootstocks?

Problems with seedling trees . Long juvenility . Thorniness . Soil-related

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Diseases

. . Diaprepes . Viruses . Blight . Nematodes

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Phytophthora (P. nicotianae, P. palmivora)

Damping-off of seedlings, foot rot, crown rot, fibrous root rot.

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Diaprepes root weevil (Diaprepes abbreviatus)

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Phytophthora Diaprepes (PD) complex

Root injury due to feeding of Diaprepes larvae predisposes citrus roots to Phytopthora infection.

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Abiotic factors

. Cold/Frost . Drought . Flooding . Salinity . pH

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Horticultural traits

. Tree size . Yield . Fruit quality

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Rootstock effect on tree size

Low vigor

Three-year-old Valencia trees

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Rootstock effect on tree size High vigor

Three year-old Valencia trees

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Requirements of a good rootstock . Reduction of juvenility. . Induce high fruit yield. . Enhance fruit quality of the scion (size, shape, brix, acids). . Adaptation to different soil types. . Resistance to abiotic stresses (salinity, drought, flooding, pH). . Resistance to pests and diseases.

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Rootstock testing (short-term)

Greenhouse screening for disease tolerance.

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Rootstock testing (long-term)

Field trials in commercial settings to assess tree growth and size, compatibility, time to bearing, etc.

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Rootstock testing (long-term)

Field trials to assessing fruit quality and yield.

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Other requirements . Compatibility with the scion. . Combination must be long-lived. . Ease of propagation in the nursery. . Should produce many fruits and seeds. . High degree of nucellar polyembryony.

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IV. HISTORY OF ROOTSTOCKS

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Origin of grafting . Origin of grafting is uncertain. . Budding is said to have been practiced in China before A.D. . Budding or grafting of citrus was already common in the Mediterranean area in the 5th century and standard practice in the 16th and 17th centuries ().

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Origin of grafting

. In Spain and Italy, mostly seedling trees were planted in the 18th century. . Advent of Phytophthora foot rot (gummosis) forced European growers to bud on resistant rootstocks. . The universally favored rootstock was sour orange.

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Florida . Florida’s citrus industry grew out of 16th Century Spanish introductions of plants and seeds of sour orange, , , and . . Many wild “groves” became established. . Citrus was spread further by Indians and pioneers who settled in north Florida and on the east coast. . There was limited commercial cultivation. . Once better transportation was available, interest in improving horticultural practices began.

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Florida . Budded trees came into use around 1830, when wild groves of sour orange were topworked with sweet orange (Citrus sinensis). . Goal was to reduce losses from foot rot, but also to modify tree growth and improve production. . Sour orange was the favored rootstock. . Large-scale nursery propagations of budded trees started in the second half of the 19th century. . In California, foot rot was not much of a problem and the use of budded trees instead of seedling trees was debated longer.

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Sour orange Citrus aurantium . Good cropping and good fruit quality. . Cold hardy. . Tolerant to Phytophthora foot rot. . Tolerant of blight. . Susceptible to citrus nematode and burrowing nematode. . Adaptable to a wide range of different soil conditions (highly tolerant of calcareous soils). . But…Highly susceptible to the (CTV) in combination with most scions.

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New problems . CTV → stem ping and quick decline of sour orange in combination with most scions except .

Brown citrus aphid Cleopatra mandarin Sweet orange . Rough lemon dominant until the 1970s. . lime dominant in Brazil (drought tolerance).

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Cleopatra mandarin /reticulata

. Produces large trees. . Small fruit of good quality. . Can be slow to come into production. . Tolerant to CTV. . Cold tolerant. . Salt tolerant. . Adaptable to a wide range of soils. . Suitable for , , oranges, and .

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Rough lemon Citrus jambhiri . Produces large trees. . High yield and large low-quality fruit. . Suitable for orange and grapefruit. . Poor cold hardiness. . Requires less irrigation. . Susceptible to wet soils. Best on deep sandy soils. . Susceptible to foot rot. . Tolerant to CTV. . But…Highly susceptible to blight (young tree decline).

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More problems

. Blight (young tree decline) first described in 1894. Prevalence increased in the 1970s with the decline of sour orange and use of rough lemon. → Trifoliate orange rootstocks Carrizo citrange Troyer citrange Swingle citrumelo

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Trifoliate orange Poncirus trifoliata . Produces small trees. . High yield and high-quality fruit. . Used mostly for mandarins, oranges, and . . Good cold hardiness. . Susceptible to highly calcareous soils. . Sensitive to high salt. . Tolerant to foot rot. . Tolerant to CTV. . Highly susceptible to blight.

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Carrizo citrange C. sinensis x P. trifoliata . Produces standard size trees. . High yield and large good-quality fruit. . Moderately cold tolerant. . Tolerant to foot rot. . Low salt tolerance. . Resistant to burrowing nematode. . Adapted to a wide range of soils, except highly calcareous soils. . Tolerant to CTV. . But … Susceptible to blight

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Swingle citrumelo C. paradisi x P. trifoliata

. Produced in 1907, but not released until 1974. . Cold tolerant. . Trees vary in vigor depending on the scion. Valencia trees are average. Grapefruit trees are larger. . Not suitable for heavy clay (flatwoods) or highly calcareous soils. . Moderately salt and drought tolerant. . Resistant to the citrus nematode. . Tolerant to CTV and blight. Walter T. Swingle

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1953-1974 Sour orange

1974-1988 Rough lemon

Carrizo 1988-2015

Swingle

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Modern rootstocks UF/IFAS breeding program UFR-1, UFR-2, UFR-3, UFR-4, UFR-5, UFR-6, UFR-7, UFR-8, UFR-9, UFR-10, UFR-11, UFR-12, UFR-14, UFR-15, UFR-16, UFR-17, …

USDA breeding program US-802, US-812, US-897, US-942, US-1279, US-1281, US-1282, US-1283, US-1284, US-1516, SS1, SS2, SS3, …

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https://www.freshfromflorida.com/

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2017-2018

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V. ROOTSTOCKS AND HLB

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Huanglongbing (HLB)

Asian citrus psyllid

InfectedHealthy Candidatus Liberibacter asiaticus (CLas)

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Definitions

Pagán and García-Arenal (Int J. Mol. Sci. 2018)

The two major mechanisms of plant defense against pathogens are: 1) resistance (the host’s ability to limit pathogen multiplication) and 2) tolerance (the host’s ability to reduce the effect of infection on its fitness regardless of the level of pathogen multiplication).

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HLB field tolerance of rootstock trees

US-897 US-942

Citrus reticulata × Poncirus trifoliata

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Greenhouse studies

Control Infected ControlInfected Control Infected

Cleopatra US-897 US-942

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Grafting

HLB-susceptible!

Grafting removes many rootstock-specific leaf biochemical compounds that may be associated with HLB tolerance.

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How do we define rootstock tolerance in a commercial grafted tree? The ability to maintain tree health and productivity despite infection. This is best measured relative to some other “standard” rootstock under the same conditions.

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Field observations . None of the rootstock prevent a grafted susceptible scion from becoming infected with CLas. . But some rootstocks enable the grafted tree to better cope with HLB. WHY?

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Metabolomics Relative concentration Relative concentration

The rootstock can influence the scion metabolically

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Rootstock can influence the scion metabolic response to HLB

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Reciprocal influence of rootstock and scion challenged with CLas

Albrecht and Bowman (2019) Scientia Horticulturae 254

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Disease progression

Scion/rootstock combination 2 mai 4 mai 6 mai 9 mai 12 mai Cleopatra/US-802 1.0 2.4 2.8 2.0 3.4 Cleopatra/US-897 1.0 1.8 2.1 1.8 2.8 Cleopatra/US-942 1.0 1.8 2.1 2.4 3.1 Cleopatra/Cleopatra 1.0 2.3 2.6 2.3 3.0 US-802/Cleopatra 1.0 1.0 1.0 1.1 1.3 US-897/Cleopatra 1.0 1.0 1.1 1.0 1.0 US-942/Cleopatra 1.0 1.0 1.0 1.0 1.1 US-802/802 1.0 1.0 1.1 1.0 1.1 US-897/US-897 1.0 1.0 1.1 1.0 1.3 US-942/US-942 1.0 1.0 1.2 1.1 1.1 Valencia/US-802 1.0 1.4 2.2 2.5 3.3 Valencia/Cleopatra 1.0 1.0 1.7 2.5 2.7 Trees with a tolerant in the scion position were less damaged by CLas infection and became infected at a lower rate than trees with a susceptible cultivar in the scion position.

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What does this mean?

. Results indicates a greater influence of the scion cultivar than rootstock cultivar on tree tolerance to infection with CLas.

. This does NOT indicate the rootstock plays no role in tree tolerance to HLB!

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Other factors . Rootstock vigor. . Better ability to cope with other soil-borne stresses and diseases under the weakened condition caused by HLB.

. Rootstock-specific architectural and anatomical traits that are associated with efficient uptake and transport of nutrients and water.

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Root structural differences

Swingle US-897 US-942

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Summary  Rootstock influences a grafted citrus tree in many ways and is very important for the success of a citrus operation.  The success of a rootstock depends on the genetic potential and on the interaction with climate, soil, pests and diseases, and cultural practices.  Rootstocks can be used to reduce the negative impacts of HLB. Even when a rootstock is highly tolerant to a specific stress or disease, it will not thrive in the presence of unfavorable conditions unrelated to that specific stress or disease.

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Ute Albrecht [email protected]

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