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Por Que Investigar El Injerto en Plantas
Por Que Investigar El Injerto en Plantas
Por Que Investigar El Injerto en Plantas
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Francisco Pérez-Alfocea
Spanish National Research Council
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Keywords: food security, sustainable agriculture, fruit quality, new cultivars, R×S×E,
new applications, transgrafting, societal and environmental challenge
Abstract
Agricultural productivity must increase by 60% to feed the expected
population of 9.6 billion people in 2050, and must be achieved through a socio-
environmental sustainability of the natural resources. However, vegetable
production around the world is being increasingly hampered by the unfavourable
soil and environmental conditions that include abiotic constraints such as drought,
extreme temperature, salinity, flooding, low nutrients, organic and heavy metals
contamination, as well as biotic ones assoil and air-borne pests and diseases. This
situation is aggravated by successive cropping, environmental policies, as the phase-
out of the chemical soil disinfectants, and negative impacts of climate change. While
genetic breeding cannot always provide efficient solutions, vegetable grafting has
developed very quickly in the last 50 years, mainly to induce shoot vigor and to
overcome soil borne diseases in susceptible crops, as a surgical alternative to
breeding. However, both the development of new rootstocks and new applications in
agriculture for crop improvement are hampered by the limited knowledge existing
on the physiological and genetic mechanisms underlying rootstock × scion
interactions × environment interactions. A major goal of research in vegetable
grafting should be gaining sound understanding about the physiological and genetic
basis of those interactions and the agronomic performance of rootstock-mediated
traits in order to establish a solid scientific basis for developing efficient rootstocks,
grafting technologies and management practices for different horticultural species
and specific environments. Scientific and technical collaboration will further
contribute to a wider development and exploitation of vegetable grafting towards
socio-economic-environmental sustainable agriculture and food security.
INTRODUCTION
Even though the benefits of using grafted seedlings are now fully recognized
worldwide, it has long been considered as an empirical process and more as a craft secret
than science. The need to understand the scientific basis of grafting-mediated crop
improvement has prompted in the most recent two decades many scientific actions aiming
at filling the gap between empiricism and science. The number of scientific publications
using grafting as an approach has drastically increased since 1992 (see Colla (2010) for
details), in accordance with the number of grafted seedlings produced worldwide.
Nowadays, many scientists employ grafting between different vegetables to (1) improve
understanding of how root-to-shoot signalling alters scion physiology and phenotype; (2)
study specifically how the root system responds to different stresses, and how these
responses affect plant adaptation; (3) gain expertise in using grafting to solve phyto-
pathological and agronomic problems such as soil-borne diseases or abiotic constraints;
and (4) alter positively fruit quality parameters.
Important efforts are being made through different international initiatives to
compile this scattered pre-existing knowledge in order to build-up a global vegetable-
a
alfocea@cebas.csic.es
grafting community and to link this knowledge to new genetic, physiological and
agronomic information, thereby contributing to better selection and exploitation of the
available genetic variability in rootstocks, and generation of new variability through
genetic and biotechnological breeding approaches. Several comprehensive reviews on
grafting have compiled existing information about historical aspects and the current state
of the art in herbaceous vegetable and ornamental plants, including practical and
agronomical aspects such as implementation, rootstocks, species and crop performance
(Lee and Oda, 2010). Other studies are partially focused on very specific aspects of root-
to-shoot communication (hydraulics and hormones) in vegetables (Ghanem et al., 2011;
Pérez-Alfocea et al., 2010, 2011), genetics and breeding (King et al., 2010), response of
grafted plants to salinity (Colla et al., 2010), nutrient and heavy metals stresses (Savvas et
al., 2010), thermal stress and organic pollutants (Schwarz et al., 2010), pathogens (Guan
et al., 2012; Louws et al., 2010) and fruit quality (Rouphael et al., 2010). Based on the
available data on the literature, it can be stated that the use of adequate rootstocks is an
efficient strategy to increase the yield stability and sustainability of the intensive
horticulture, and that it has an enormous potential for contributing to food and
environmental security. As American farmers say, “there’s not only a market for it,
there’s a great need for it”, where the ‘need’ not only implies benefits for agriculture, but
indirectly also for the environment and the society.
Rootstocks can contribute to increasing yield stability and sustainability in
vegetable crops through (i) increasing the yield potential of elite cultivars; (ii) filling the
yield gap under biotic (i.e. fungi, bacteria, virus, nematodes) and abiotic (i.e. salinity,
drought, low nutrients, extreme temperatures, soil compaction) stress suboptimal
conditions; (iii) decreasing the use of chemical (pesticides and fertilizers) and
contamination in soil and aquifers; (iv) increasing the efficiency of use of natural (water
and soil) resources; and, therefore, (v) optimizing crop productivity. The socioeconomic
and environmental potential impact of grafting is as broad as the genetic variability able
to cross the barrier between the rootstock and the scion. However, only through
understanding the mechanisms underlying the agronomical phenotypic variability
resulting from rootstock × scion × environment interactions will allow further
development and exploitation of rootstocks for increasing yield stability and sustainability
in horticultural crops. The results obtained so far are very promising, but more research
must be focus on (i) germplasm identification and rootstock breeding; (ii) rootstock-
mediated mechanisms regulating shoot growth; (iii) physiological and genetic
mechanisms involved in (a)biotic stress resistance; (iv) optimization of management
practices; (v) rootstock influence on fruit quality; (vi) generation of new (epi)genetic
variability and understating the mechanisms involved; (vii) use of biotechnological
approaches as transgrafting; and (viii) increasing grafting use efficiency.
22
variability. Indeed, it is expected that the major development of rootstocks will occur in
Solanaceae, Cucurbitaceae and other annual crops including other families such as
Fabaceae (soybean, peanut), Asteraceae (artichoke) and Euphorbiaceae (cassava). Both
Solanaceae and Cucurbitaceae include 98 and 125 genera and 2700 and 960 species,
respectively, with additional accessions and ecotypes. More than 1000 million tons of
vegetables are produced annually in the world, with around 53 million hectares cultivated
(FAOSTAT, 2010). Many of them are either grafted or can be grafted. Much more well-
performed screening for specific environments, development of hybrid combinations and
graft-compatibility studies are required.
23
sources, but requires much more extensive and intensive work. Indeed, it is difficult to
separate the rootstock-mediated vigor and resistance effects. For instance, the 25-55%
yield increases in grafted melons affected by Fusarium were not only due to the disease
resistance but also especially to plant vigor maintenance during the late growing season
(Lee et al., 2010).
Besides the vigor or genetic resistance, the rootstocks have also the capacity to
provoke changes in the rhizosphere that can positively (association with beneficial
microorganisms) or negatively (changes in pathogenic variation of existing
microorganism and/or re-emergence of new ones) affect plant performance and crop
productivity (Gilardi et al., 2013; Pérez-Alfocea et al., 2011). It should be expected that
understanding the physiological and genetic mechanisms underlying this root × beneficial
microorganism interaction will contribute to incorporate new interesting traits to the
commercial rootstocks aiming at a better yield stability and soil sustainability.
24
acids, pigments like lycopene and other carotenoids and some vitamins (Rouphael et al.,
2010). However, no attention has been paid so far to other minor organic compounds that
can be specifically produced by the new roots and transported to the fruits, which can
influence the nutritional, nutraceutical and toxicological properties of the fruits, being an
important factor to be considered in terms of food security, in addition to yield stability
under a changing environment.
A specially promising rootstock for tomato interspecific grafting is Solanum
torvum, a species native to the western tropics and India that tolerates the climatic
pressures of tropical regions (floods, drought, high temperature) and a wide range of soil
borne pathogens, including Verticillium dahlia, Ralstonia solanacearum, Fusarium
oxysporum and Meloidogyne spp. root-knot nematodes (Petran and Hoover, 2014).
However, even if S. torvum has been selected as rootstock of interest against other tomato
rootstocks of known high compatibility, this species is currently classified as ‘‘noxious
weed’’ which prohibits, without permit, its movement ‘‘into or through the United States,
or interstate’’ (ECFR, 2008), in addition to other additional restrictions that have been
placed by several US states (Smith et al., 2008).
Intoxication outbreaks have been detected by consuming 20-30 mg eq
solanine/100 g fruit of S. torvum (Smith et al., 2008). Toxic effects appear to be mediated
by several steroideal glycoalkaloids (SGAs) including solasonine and solamargine,
isomers of a chacotriosyl hydroxysolanidine and possibly other compounds which
produce gastrointestinal and neurological effects (Rangan, 2008; Smith et al., 2008).
Those toxic compounds are derived from cholesterol and can act synergistically in the
plant resistance to insects, fungi, and nematode (Morris and Lee, 1984). Although toxic
SGAs are not usually detectable in the fruit, tomato leaves and vines contain SGAs
similar to solanine, and the fruits may be poisonous under certain circumstances and it has
been speculated that pre- and post-harvest stresses, fruit maturity, cultivar differences,
and growth environment can contribute to substantial changes in final concentration into
the fruit (Smith et al., 2008). Therefore, the existence of a transport from the S. torvum-
derived rootstock to the scion cannot be ruled out and should be investigated.
Except for some chlorophyll and photosynthesis-related compounds, it is believed
that roots are able to produce every type of compounds that exist in plants (Gregory,
2006; Uren, 2001). Special attention has been paid to the study of the organic compounds
released from plant roots into the rhizosphere in relation to the soil and biota interaction,
where metabolites from major chemical groups from primary and secondary metabolism
(sugars and polysaccharides, amino acids, fatty acids, sterols, vitamins, enzymes,
flavonones, carotenoids, hormones) have been identified (Gregory, 2006; Uren, 2001).
Whether or not those compounds are present in the root-xylem sap and can be transported
and accumulated into the fruit remains an open question that need to be addressed.
25
TRANSGRAFTING
Transgrafting can be defined as the use of a genetically engineered rootstock to
support a WT scion, or vice versa, and it might be very useful to protect horticultural
crops from biotic and abiotic stresses (Lemgo et al., 2013) and can be considered as a
promising biotechnological tool to address food security, beyond conventional breeding
technologies (Lusser et al., 2012). Creating new transgenic rootstocks to resist biotic (e.g.
(fungi and viruses) and abiotic stresses (salinity, drought, soil deficiencies,
temperature,…), especially those for which resistance genes are rare in the plant genome,
is a promising development (Lusser et al., 2012). Although the presence of the trans-
genetic material in the shoot tissues may be a concern since both mRNAs and siRNAs can
cross a graft junction, the use of RNA signaling technology in modified roots represents
an excellent opportunity to induce virus resistance(s) but still maintaining production of
non-transgenic fruits. However, transgrafting might help to minimise consumer concerns
of transgene flow, since the non-transgenic scion would only produce non-transgenic
pollen (COGEM, 2006; Lev-Yadun and Sederoff, 2001). The potential of
biotechnological approaches addressed to the rootstock are obvious.
ACKNOWLEDGEMENTS
We thank the European Commission (ROOTOPOWER Contract # 289365), the
EU COST office (FA1204 Action on Vegetable Grafting), and the Spanish MICINN-
FEDER (AGL2011-27996) for support grafting research in my laboratory.
26
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Figures
50 1000
Korea
South-Eastern Spain
40 Number of publications 800
Million seedlings
30 600
Number
20 400
10 200
0 0
2000 2002 2004 2006 2008 2010 2012
Year
Fig. 2. Future challenges and prospects in vegetable grafting that should be addressed by
current and future research initiatives.
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