Fusarium wilt disease intensity of grafted and nongrafted tomato plants onto solanum rootstocks.
\r\n\tThis book shall focus on these antisense guided sequence specific silencing molecules with different mechanisms and potency for gene silencing, providing the reader with a comprehensive overview of the current state-of-the-art in ASO based therapeutics, featuring the more recent developments in terms of clinical translation and the use of nanomedicine for the effective delivery of therapeutic nucleic acids towards precision medicine.
",isbn:null,printIsbn:"979-953-307-X-X",pdfIsbn:null,doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"96f256f5bb2e750c7496b3c0b62cb95a",bookSignature:"Prof. Pedro Baptista and Prof. Alexandra R Fernandes",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/9571.jpg",keywords:"gene therapy, gene silencing, genome modulation, post-transcriptional modulation, modified oligonucleotides, PNAs, LNAs, siRNA, antisense nucleotides, vectorization of antisense nucleotides, nanotheranostics, clinical translation, nanoparticles for gene delivery",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:null,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"October 25th 2019",dateEndSecondStepPublish:"November 15th 2019",dateEndThirdStepPublish:"January 14th 2020",dateEndFourthStepPublish:"April 3rd 2020",dateEndFifthStepPublish:"June 2nd 2020",remainingDaysToSecondStep:"a year",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:null,kuFlag:!1,biosketch:null,coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"82671",title:"Prof.",name:"Pedro",middleName:null,surname:"Baptista",slug:"pedro-baptista",fullName:"Pedro Baptista",profilePictureURL:"https://mts.intechopen.com/storage/users/82671/images/system/82671.jpg",biography:"Pedro Viana Baptista (b.1972) holds a degree in Pharmaceutical Sciences (1996) from the Universidade de Lisboa. He obtained his PhD in Human Molecular Genetics from the School of Pharmacy, University of London in 2000. In 2001 moved to FCT-NOVA where he created the Nanomedicine Group, which he leads. Currently, he is Full Professor of Molecular Genetics & Nanomedicine at the Department of Life Sciences, FCT-NOVA and responsible for the NanoImunoTech Group – Nanomedicine in the Applied Biomolecular Sciences Research Unit. His work focuses on the biomedical applications of nanoparticle-based strategies towards light-induced cancer therapy and as gene silencing platforms (including siRNA, antisense and nanobeacons). Coordinates several research projects focused on the use of nanotechnology for molecular diagnostics and nanotheranostics, including nanoparticles for diagnostics and therapy; biosensors (TFTs and ISFETs); medium-throughput SNP analysis platforms, and nanoparticle-based therapies (nanovectors for siRNA and antisense therapy, targeted combined therapies).",institutionString:"Universidade Nova de Lisboa",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"2",totalChapterViews:"0",totalEditedBooks:"0",institution:{name:"Universidade Nova de Lisboa",institutionURL:null,country:{name:"Portugal"}}}],coeditorOne:{id:"253664",title:"Prof.",name:"Alexandra R",middleName:null,surname:"Fernandes",slug:"alexandra-r-fernandes",fullName:"Alexandra R Fernandes",profilePictureURL:"https://mts.intechopen.com/storage/users/253664/images/system/253664.jpg",biography:"Alexandra R. Fernandes is an Assistant Professor at the Department of Life Sciences, FCT-NOVA where she leads the group of Cancer Therapeutics dedicated to assessing novel compounds against tumor cells and elucidate the underlying molecular mechanisms. 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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"72",title:"Ionic Liquids",subtitle:"Theory, Properties, New Approaches",isOpenForSubmission:!1,hash:"d94ffa3cfa10505e3b1d676d46fcd3f5",slug:"ionic-liquids-theory-properties-new-approaches",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/72.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"314",title:"Regenerative Medicine and Tissue Engineering",subtitle:"Cells and Biomaterials",isOpenForSubmission:!1,hash:"bb67e80e480c86bb8315458012d65686",slug:"regenerative-medicine-and-tissue-engineering-cells-and-biomaterials",bookSignature:"Daniel Eberli",coverURL:"https://cdn.intechopen.com/books/images_new/314.jpg",editedByType:"Edited by",editors:[{id:"6495",title:"Dr.",name:"Daniel",surname:"Eberli",slug:"daniel-eberli",fullName:"Daniel Eberli"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"57",title:"Physics and Applications of Graphene",subtitle:"Experiments",isOpenForSubmission:!1,hash:"0e6622a71cf4f02f45bfdd5691e1189a",slug:"physics-and-applications-of-graphene-experiments",bookSignature:"Sergey Mikhailov",coverURL:"https://cdn.intechopen.com/books/images_new/57.jpg",editedByType:"Edited by",editors:[{id:"16042",title:"Dr.",name:"Sergey",surname:"Mikhailov",slug:"sergey-mikhailov",fullName:"Sergey Mikhailov"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1373",title:"Ionic Liquids",subtitle:"Applications and Perspectives",isOpenForSubmission:!1,hash:"5e9ae5ae9167cde4b344e499a792c41c",slug:"ionic-liquids-applications-and-perspectives",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/1373.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"2270",title:"Fourier Transform",subtitle:"Materials Analysis",isOpenForSubmission:!1,hash:"5e094b066da527193e878e160b4772af",slug:"fourier-transform-materials-analysis",bookSignature:"Salih Mohammed Salih",coverURL:"https://cdn.intechopen.com/books/images_new/2270.jpg",editedByType:"Edited by",editors:[{id:"111691",title:"Dr.Ing.",name:"Salih",surname:"Salih",slug:"salih-salih",fullName:"Salih Salih"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"63534",title:"Hydrogen Water on Survival Rate after Fasting in Drosophila Model",doi:"10.5772/intechopen.80777",slug:"hydrogen-water-on-survival-rate-after-fasting-in-drosophila-model",body:'\nIn recent years, there has been a great deal of attention toward the field of free radical chemistry. Our body generates free radicals of reactive oxygen species or reactive nitrogen species by various endogenous systems, exposure to different physiochemical conditions. To balance the free radicals and antioxidants is necessary for proper physiological function. If we can regulate free radicals to reduce the potential reactive oxygen species over tension on body’s ability, it will not cause oxidative stress damage. Free radicals thus do not adversely alter lipids, proteins, DNA and trigger many diseases. As a result application of an external source of antioxidants likes oral hydrogen water can assist in coping this oxidative stress.
\nThis metabolism phenomenon also occurs in many other animals or even insects that are less evolved. In 1946, the fruit flies were successfully carried out by National Aeronautics and Space Administration (NASA) for the biological experiments in space. About 75% of the genetic code of fruit flies has similarities to all human pathogenic genes. Insect life is much shorter than human. Drosophila flies also has found free radicals in their cells. The scientists take Drosophila to study aging effects much efficiently due to shortening the time in experiments. It is an important reason why studying Drosophila melanogaster in fruit flies is very popular among biological researchers. They also found many mutant species of fruit flies, which are related to the regulatory function of oxidative stress attacked by free radicals in cells. Therefore, we study the survival effects of drinking hydrogen-water on fruit flies after fasting.
\nAccording to past research reports, fasting or severely hungry animals will rapidly accumulate a large number of free radicals in cells because of lack of well physiological functions. Excessive free radicals, however, can damage the cell’s genetic material and cell membranes, eventually killing the fruit fly. And the flies of the mutant species are more likely to accumulate free radicals in vivo because the chromosome of the Drosophila cell, an enzyme gene that removes peroxidase free radicals by hydrazine, is removed so that the number of free radicals is out of control. Although the fruit fly of this mutant species can still develop to adults, its lifespan is shorter than that of wild fruit flies. I used this kind of fruit fly to do experiments because they have accumulated many free radicals in the body, and alkaline hydrogen water (negatively charged) can neutralize positively charged free radicals, so feeding hydrogen-water on the mutant flies are more evident than wild species. In the course of the experiment, I found that the mutant flies fasting (stop feeding water), causing the flies to die. It proved that the fruit fly did have poor anti oxidative stress, so I did not fast-feed the fruit flies in advance during the experiment. Only for the wild flies, fasting beforehand.
\nIn the study, the experimental result shows that the mutant fruit flies of Drosophila melanogaster have a much stronger response to hydrogen water than wild ones. It appears to support a possibility that hydrogen-water seems to be a definite help for individuals who are more likely to accumulate free radicals. Therefore, it is the best way to use the mutant flies directly for anti-free radicals experiments. There will be more significant results. It is also possible to test older flies that are more sensitive to alkaline electrolyzed water because the larger the flies, the more free radicals will accumulate in the body. In our experiments, we found that the older the flies, the more they can’t afford the pressure of fasting. On the other hand, the open trial can also directly use strong alkaline hydrogen water to feed mutant flies. In addition, the moisture of the feeding environment in the test tube during the experiment is also vital; otherwise, the fruit flies will die a lot and affect the experimental result. In the same operation, the problem of escaping of the flies is also to avoid. Otherwise, if the trial flies are too few, they are not representative.
\nThe untreated tap water is almost neutral at about pH 7.0 with a positive oxidative potential of +150 mV. The tap water after filtered and electrolysis treated, the cathode hydrogen dissolving water belongs to be alkaline at a pH over 8.0 with a relatively high negative reductive potential of −150 to −300 mV. Recently, a lot of literatures [1, 2, 3] found that drinking hydrogen water increases the activity of a critical detoxifying enzyme of superoxide dismutase in the body. It protects against free radicals toxic damage on protein, enzyme, DNA, lipid, and membrane in cells (see Figure 1). Since the alkaline hydrogen water molecules with an extra amount of free electrons, which can neutralize the highly reactive free radicals before free radicals take away free electrons from intracellular molecules.
\nFree radicals toxic damage on protein, enzyme, DNA, lipid, and membrane.
Water, which constitutes over 70% of the body, is involved in virtually every function of life. It forms the bodily fluids, such as blood, lymph, cerebrospinal fluid, saliva, and digestive fluids to regulate the metabolism of joint lubrication, detoxification, and maintaining the blood pressure. While the water molecules around the cell membranes, they show a long-range ordering feature and like an epitaxial liquid crystal with distinct properties from the bulk state [1]. Shirahata et al. [2] showed that drinking hydrogen water increases the activity of a critical detoxifying enzyme, superoxide dismutase (SOD), which exists in humans, animals, plants, and micro-organisms, as an essential antioxidant to protect cell under oxidative stress damage in vitro. Superoxide is one of the primary reactive oxygen species in the cell. So SODs serves a key antioxidant role. This antioxidant function exerted by molecular hydrogen has been proved [3]. In the rat model, drinking hydrogen water showed signs in the decrease of the peroxided lipid level in their urine. When comparing to tap water to hydrogen-water, the rat drinks hydrogen-water can extended to live 20–50% longer [4]. Finally, we studied the survival rate of drinking hydrogen-water on Drosophila flies after fasting, which related to the regulatory function of oxidative stress attacked by free radicals in cells.
\nDrosophila lacking SOD1 has a dramatically shortened lifespan, whereas fly lacking SOD2 will die before birth. Lacking SOD3 does not show any visible defects and exhibit an average lifespan, though more sensitive to hyper oxidative injury. Drosophila melanogaster is a species of fruit flies. Drosophila fly is typically used in research because it can readily rear in the laboratory. Since Charles W. Woodworth used Drosophila melanogaster as modal organism, eight Nobel prizes have awarded to research using Drosophila. It continues to be widely applied to biological research in genetics, physiology, microbial pathogenesis, and lifespan [5]. Wild-type fruit flies are yellow-brown, with brick-red eyes and transverse black rings across the abdomen. Females are about 2.5 mm long, and males are slightly smaller with darker backs (see Figure 2).
\nFemale (left) and male (right) of Drosophila melanogaster [6].
The wild-type and mutant-type flies obtained from the Drosophila melanogaster stock center at the National Health Research Institutes in Taiwan. Peroxide reductase-1 is one of the mutant-type fruit flies. Drosophila melanogaster flies have every six tubes of females and males of 2 day-old-adult flies, 30 flies per tube. Four ones (20/tube) of females and four ones (30 tubes/tube) of males collect within 2 day-olds of adult flies. These flies gently placed to the bottom of the vial (9.5 cm tall × 2.4 cm diameter) and top with cotton swabs and balls with pure neutral water and alkaline hydrogen water (pH 8.5 and 9.5) in the standard medium supplement at room temperature of 50–70% relative humidity. Before experiments, each group carefully collected 2 day-old males and females of adult flies, respectively. The experimental procedures are as follows: 1. Wild-type flies have fasted for 12 hours and then feed water, while the munt flies do not. 2. Using wetted cotton swabs inserted into the test tube for two feeds a day. 3. Observe the death of the flies every 12 hours until all the flies have died. 4. All experiments treatments kept at the standard medium. 5. If the feeding tube wetted with water droplets and change it periodically.
\nFor the control group, every two tubes are wild and mutant species of fruit flies feed with pure water of pH 7. One is female fly, and another is male fly. For the treatment group, every four tubes are wild and mutant species with the female and the male ones fed with alkaline hydrogen water of pH 8.5 and 9.5, respectively. Generally, regardless of whether it is a wild fruit fly or a mutant fruit fly, in the fasting state, feeding of hydrogen water has a significant increase in the survival rate (see Figures 3, 4, 5, 6).
\nSurvival percent in % of female-mutant fruit flies vs. day after fasting. Note: pH 7: pure water; pH 8.5: weak-alkaline-H2-water.
Survival percent in % of male-mutant fruit flies vs. day after fasting. Note: pH 7: pure water; pH 8.5: weak-alkaline-H2-water.
Survival percent in % of female-wild flies vs. day after fasting. Note: pH 7: pure water; pH 8.5: weak-alkaline-H2-water; pH 9.5: medium-alkaline-H2-water.
Survival percent in % of male-wild flies vs. day after fasting. Note: pH 7: pure water; pH 8.5: weak-alkaline-H2-water; pH 9.5: medium-alkaline-H2-water.
If we compare with the results of the mutant and wild species, the mutant fruit fly of males to feed weak alkaline hydrogen water of pH 8.5 is the most significant change in survival among them (see Figure 3 vs. Figure 5; Figure 4 vs. Figure 6). In the case of feeding mutant flies with pure water of pH 7, for example, when fasting for more than 4 days, half number of deaths occur. If weak alkaline hydrogen water of pH 8.5 fed, it will delay to more than 9 days to happen it (see Figure 3). That is about a twice of increase in the lifespan. Therefore, a weak alkaline hydrogen water of pH 8.5 seems to have a positive effect on the survival rate of flies in the state of starvation.
\nPast research results have shown that the average lifespan of females is longer than that of males. Our experimental results showed that the effect in response to the feeding of hydrogen water, the male-fly was significantly more than the female-fly for all the wild and mutant species. While the case of feeding wild flies with pure water, for example, fasting for more than four to 6 days, to dead a half (see Figures 5 and 6). If hydrogen water supplied, fasting time of female fly and male fly extended for more than 7 days. However, hydrogen water seems to have a less help on the survival rate of wild female flies in the state of starvation (see Figure 5). This result implies that hydrogen water appears to be of much more help to individuals with weaker constitutions than to individuals who have stronger ones.
\nTested for the relationship between the alkaline pH (pH 8.5 and 9.5) and the survival response. Our previous results have shown that drinking hydrogen water seems to be helpful for the survival rate of Drosophila in the case of starvation. Therefore, it is worthwhile to analyze hydrogen water in alkalinity further the impact on the survival rate of Drosophila. When the pH of the hydrogen water increased from 8.5 to 9.5, after fasting time more than 4 days the significant increase in the survival rate of the male fly observed in Figure 6. However, for the female fly, the result is not the same. Interestingly, although the wild species of female fruit fly, hydrogen water increased from pH 8.5 to 9.5, the survival response was slightly decreased. That is to say, further addition in alkalinity, however, has the opposite effect on the female flies, and the survival rate does not rise any longer.
\nIn the past, many works of literature have reported that human aging may be related to the function of free radicals to destroy cells. However, why do free radicals come? For example, the role of cells depends on the oxidation and decomposes the nutrients in the body. It is like gasoline, which burns oxygen to release heat energy and promote steam. However, in the process of redox, cells produce a byproduct, oxygen free radicals. The oxygen molecule (O2) itself has 16 electrons, but it loses one electron in the process of the redox reaction, and the oxygen that becomes a single electron becomes very unstable, and it will destroy the function of the cell and even cause disease. Interestingly, if the neutral water is decomposed using a water electrolysis generator, the water molecules will change to negatively charged alkaline hydrogen water. If the electrolytic liquids are separated, acidic water will act like oxygen free radicals to destroy the cells. However, hydrogen water is the opposite, with an extra free electron, which can combine with free radicals to neutralize the ability to equilibrate free radicals before free radicals take away an electron from intracellular molecules. Therefore, hydrogen water may have anti-oxidant function and protect cells from free radical damage.
\nThis experiment supports that drink hydrogen water may help the fasting fly survival rate. When oxygen free radicals in the intestine are out of balance, hydrogen water may neutralize free radicals and reduce damage to cells. Drosophila has an innate immune system similar to humans, and the structure and physiology of the fruit fly’s gastro-intestine, cardiac, and neurological diseases resemble that of humans. Even if the genes and mechanisms that they are involved whether conserved or not, the biological process in a similar genes species often provides a valuable framework to study anti-oxidative stress effect and allow development of potential clinical applications.
\nWhen Drosophila intestinal oxygen free radicals increase due to fasting, the increased reactive oxygen species in the intestine pass through nitric oxide, erythrocytes, and another non-nitric oxide signal to activate transcription of NF-κB protein in the fat of Drosophila. The factor, Drosophila liver, promptly initiated in the response of cells to many stimuli, including oxidative stress, cytokines, free radicals, ultraviolet radiation, and immune response of the antibacterial peptide, causing a systemic immune response in the Drosophila. The biological model of Drosophila used to simulate that the immune response of human organs is closely related to intestinal health [7].
\nHowever, this does not mean that drinking hydrogen water is beneficial to all healthy individual because, in the body, some enzymes that regulate free radicals, which can balance the problems caused by the accumulation of free radicals. Therefore, drinking a significant amount of hydrogen water in a healthy state may be interference to the body’s natural regulation mechanism. In other words, as healthy people do not need and should not take medicine, only those who are sick need to follow the doctor’s instructions. All in all, the function of hydrogen water on the human body needs further experiments to be further studied. This experiment can only support the flies in the extreme unfavorable environmental pressure; feeding hydrogen water may increase its survival rate.
\nGrafting is the deliberate joining together of a scion and rootstock, taken from different but compatible plants, which are taxonomically close, to produce a composite plant. The scion, which forms the top portion, is selected for its desirable attributes, such as better yields, bigger fruit sizes, or preferred flavor. The rootstock onto which the scion is grafted is selected for reasons such as its vigorous growth and resistance/tolerance to soilborne diseases and pathogens as well as its ability to withstand soil extremes [1]. The technique of grafting vegetables originated from Japan and Korea in the late 1920s. The first record of an interspecific graft for increased yield and pest and disease control was reported in Japan between watermelons [Citrullus lanatus (Thunb.) Matsum and Nakai] as scion and squash (Cucurbita moschata Duch.) The watermelon grafting technique was then widely introduced to farmers in Japan and Korea between the 1920s and 1930s; later, the technique was extended to grafting of other vegetable crops Cucumis sativus L. [2] and Solanum melongena L. in the 1950s [2] and then to Lycopersicon esculentum Mill [1].
Vegetable grafting is implored to impart resistance to soilborne pathogens, for example, nematodes [1, 3] and increase yields [3] and tolerance to abiotic stress conditions [4, 5, 6, 7, 8].
Tolerance to soilborne diseases is one of the main reasons why vegetable grafting is practiced. Rootstocks are selected based on their tolerance to common vegetable production diseases caused by Verticillium, Phytophthora, Fusarium, and nematodes [3, 9, 10, 11].
Vegetable grafting has been shown to increase fruit yields of vegetables such as tomato and eggplants and enhances nutrient uptake together with improved water use efficiency [3, 12]. An improved water use efficiency and nutrient uptake enables grafted plants to withstand short dry spells and also increase photosynthetic activity. Eggplant rootstocks have the ability to withstand flooding conditions for several days [13].
There are cost implications in any grafting venture, and these must be properly considered before beginning a grafting project. A positive or negative net return is mainly dependent on the cost of producing the grafted plants and the prevailing market price for the tomato fruits that will be produced [14]. Falling tomato prices coupled with high input cost for raw materials needed for grafting may result in some negative net returns. The net returns are also sensitive to the vigorousness of the rootstock and that the higher the marketable fruits, the higher the net returns. Costs of grafted plants (including seed, labor, and cost of other materials) have been estimated as $0.78 per grafted plant for 1000 plants per season in a small nursery [15]. Other investigators have also estimated the production costs of grafted and non-grafted seedlings at $0.67 and $0.15 per plant, respectively, in the production of fresh market tomato in Florida, USA [14].
Generally, labor cost represents a small proportion of the total cost of grafting, and the majority of the cost goes into the purchase of root stock seeds that are specially bred and forms 36% of the total cost [16]. However, apart from the cost of seeds, other inputs such as grafting clips and building a humidity chamber serve as additional cost.
Grafted transplants are more expensive to produce per plant than nongrafted plants. Therefore, a lower cost of rootstock can easily boost the rate at which farmers adopt this technology [15].
Grafting a selected crop variety on to another is based on the genetic attributes of both crop varieties. Farmers select rootstocks with desirable genetic properties, for example, resistance to nematodes, flooding, salinity, extreme temperatures, and increased yield production. Tomato and eggplants are the most grafted plants in the Solanaceous family, although crops of the cucurbitaceous family (melon) are also utilized [17].
The most common rootstocks used for commercial tomato grafting are hybrids (F1) or inter-specific hybrids, which have been specifically bred for resistance against pathogens and other diseases such as nematodes, Verticillium wilt, and Fusarium wilt. Hybrids are produced by crossing selected tomato varieties with other wild Solanum species with the genetic ability to offer resistance to specific diseases and pathogen infection [18].
In Europe, tomato hybrids are used as rootstocks compared to other Solanum spp., because of their high level of genetic improvements [17]. There are other plants that share the same family with tomato (Solanum torvum, S. aethiopicum, and S. macrocarpon); these can serve as rootstocks for their tolerance to waterlogged and drought conditions, Fusarium wilt, and root knot nematode infestation [13]. Most eggplant lines utilized will graft successfully with tomato lines. Rootstocks selected should be resistant to bacterial wilt (caused by, for example, Ralstonia solanacearum) and other soilborne diseases. The Asian Vegetable Research and Development Centre (AVRDC) recommends eggplant accessions EG195 and EG203, which are resistant to flooding, bacterial wilt, root-knot nematode (Meloidogyne incognita), tomato Fusarium wilt (caused by Fusarium oxysporum f.sp. lycopersici), and southern blight (caused by Sclerotium rolfsii) [13]. Grafting of a tomato variety “Pectomec” onto S. aethiopicum and S. macrocarpon in the University of Ghana Farm, Legon provided resistance to Fusarium wilt caused by Fusarium oxysporum; however, nongrafted tomato plants had a disease intensity of 46% (Table 1) and were highly diseased [19] (Figure 1). Grafting success of the tomato variety “Pectomec” onto S. aethiopicum, S. lycopersicon ”Mongal F1,” and S. macrocarpon was poor with the rootstock S. lycopersicon ”Mongal F1” [19] (Table 2).
Treatment | NRP | NDRP | DI (%) |
---|---|---|---|
Control | 24 | 11 | 46 |
P/SM | 24 | 0 | 0 |
P/SA | 24 | 0 | 0 |
Fusarium wilt disease intensity of grafted and nongrafted tomato plants onto solanum rootstocks.
NRP = Number of recording plants; NDRP = Number of diseased recorded plants; DI = Disease intensity (%); P/SA = Pectomech grafted onto Solanum aethiopicum; P/SM = Pectomech grafted Solanum macrocarpon. Agyeman [19].
Symptoms of fusarium wilt disease of tomato variety “Pectomech”. (A) Advanced symptoms (browning and wilting of leaves—red arrow), (B) Browning of tomato vascular tissues (red arrow).
Rootstocks | Number of grafted plants | Graft success | Percentage (%) graft success |
---|---|---|---|
P/M | 196 | 2 | 1 |
P/SM | 196 | 184 | 94 |
P/SA | 196 | 185 | 94 |
Grafting success of Pectomech onto three Solanum rootstocks.
P/M = Pectomech grafted onto Solanum lycopersicon ”Mongal F1”; P/SA = Pectomech grafted onto Solanum aethiopicum; P/SM = Pectomech grafted onto Solanum macrocarpon. Agyeman [19].
An ideal rootstock for tomato grafting should not only be resistant to pathogens, but also have high compatibility with the scion of tomato, with the ability to express a high level of vigorousness and resistance to pest and diseases. Rootstocks with very high levels of vigorousness compared to the scion may result in the tomato grafts being more vegetative with less fruit yield and quality [20]. Rootstocks selected should be resistant to bacterial wilt and other soilborne diseases. The tomato line (Hawaii 7996) has a high level of resistance to bacterial wilt and Fusarium wilt and is a recommended variety by AVRDC [13].
In developing countries, the use of tomato hybrids as rootstocks is limited because of the costs of imported hybrid seeds. Therefore, the use of eggplants as rootstocks is the most common method, of choice with S. torvum, S. macrocarpon, and S. aethiopicum being the most selected eggplants [21]. In rootstock selection, the eggplants are exposed to the biotic agent in pot or field evaluations, and tolerant or resistant rootstocks are selected for grafting experiments.
In a grafting study by Owusu et al. [22], against root-knot nematodes, five tomato cultivars were selected with “Big Beef,” “Celebrity,” and “Jetsetter” being resistant to Verticillium wilt, Fusarium wilt, nematodes, and tobacco mosaic virus (VFNT), which served as the nematode-resistant rootstocks, and “Tropimech” (VF) and “Power” (locally grown nematode-susceptible cultivar) served as scions. Grafted plants had the least nematode populations in the plant house. In field experiments, nematode population levels were lower in “Power” that had been grafted on Celebrity, Jetsetter, and Big Beef rootstocks, compared to self-grafted or ungrafted “Power”. Fruit yields were also higher in the grafted plants utilizing resistant rootstocks than nongrafted plants.
In another study, grafting for root-knot nematode management in heirloom tomato production was undertaken. Susceptible heirloom tomato cultivars (S. lycopersicum “Brandywine” and S. lycopersicum “Flamme”) were grafted onto two hybrid rootstocks (S. lycopersicum “Multifort” and S. lycopersicum “Survivor”); the non-grafted and self-grafted plants served as controls. Results revealed that root damage or galling significantly reduced by 81% in the hybrid rootstocks, compared to the controls. There were, however, no clear correlations between root galling and total tomato yields [15].
Tomato grafting onto a resistant rootstock of wild brinjal (S. sisymbriifolium) under farmers’ field conditions at Hemza of Kaski district against root-knot nematodes was undertaken. The root system of the grafted plants was free from gall formations; however, nongrafted plants had an average of 7.5 gall index (GI). Fruit yields significantly (P > 0.05) increased by 37% in the grafted plants compared with the nongrafted plants [23]. Eight wild Solanum rootstocks and two tomato hybrids were screened against root-knot nematode infection. Results revealed that the S. sisymbriifolium, Physalis peruviana, and S. torvum had the least galls per 10 g root (6, 5, 5) and females per g root (2, 2, 2), respectively, and showed the highest level of expression of phenolics and defense-related enzymes viz., peroxidases, polyphenol oxidases, phenylalanine ammonia lyase, and acid phosphatase from leaf samples, compared to the susceptible tomato scion (US-618) [24]. In a previous study, two garden egg rootstocks S. torvum and S. aethiopicum were poor hosts of M. javanica and M. incognita [25].
A successful grafting technique is one that would unite the scion and rootstock and enable both sections to grow together as a composite plant. The scion could be a small piece of shoot with several buds or a single bud that has been removed from an existing plant. The rootstock on the other hand forms the lower portion of the graft that forms the plant’s root system.
Several grafting techniques are used by farmers for various tree crops and vegetable production generally. In grafting of vegetables, methods such as the splice, whip and tongue, hole insertion, and pin and cleft grafting methods can be used. However, the splice/tube grafting and cleft/wedge grafting are most commonly used because of the relative ease and strong vascular connection formed between scion and rootstock. It can also be used on seedlings with age ranging from 3 to 4 weeks [26].
With the splice grafting method, slanting cuts are made on both the scion and the rootstock at an angle of 45°, and the cut surfaces are then joined together to ensure the cambium layers of the scion and the rootstock, which are properly aligned. The joined surfaces are held firmly in place with the help of a grafting clip or tube.
The cleft graft method on the other hand, involves making a clean horizontal cut on the rootstock 5 mm below the cotyledon; a 4-mm vertical incision is then made in the middle of the root stock. The scion is then sharpened in the form of a wedge and gently inserted into the incision made in the rootstock.
The selection of a particular grafting method or technique depends on the skill of the person carrying-out the grafting and the ease with which the technique can be carried out. Other factors such as the type of vegetable crop and the sowing period of the rootstock and the scion are also considered. For instance, some farmers prefer using the whip and tongue technique when grafting cucumbers because the seedlings of cucumber are large (hypocotyl length and diameter), making the grafting process easy [27].
The tube grafting method also has a high percent graft rate. The grafting of two tomato cultivars (“PG3” and “Beaufort”) using the tube and the cleft graft methods resulted in a high-percentage graft rate (79–100%), an indication of the suitability of both methods for tomato grafting [28].
There are five requirements critical to achieve a successful graft union: (1) the scion and rootstock should be compatible, (2) proper cambial alignment between scion and rootstock, (3) enough pressure to keep the cut surfaces firmly together, (4) avoidance of desiccation by maintaining high humidity around the cut surface, and lastly (5) both plants should be at the proper physiological stage for grafting to occur [29]. Good craftsmanship is an important requirement that brings the five requirements together. Graft union formation in compatible species involves a number of stages. In the first stage, parenchymatous cells are formed on the cut surfaces of the scion and rootstock followed by the interlocking of the callus between scion and rootstock leading to the formation of a callus bridge. This is followed by the differentiation of cells and the formation of the vascular cambium across the callus bridge between the scion and the rootstock and the eventual connection between phloem and xylem of the scion and rootstock to form a composite plant. The vascular connection lays the foundation for the transport of nutrients and water [30]. In tomato grafting, the formation of the xylem and phloem vessels occurs 8 days after grafting is performed [31].
Graft incompatibility refers to the inability of a graft union to form or grow properly between a scion and a rootstock, because of certain physical or chemical characteristics of the scion and rootstock. This leads to major setbacks in grafting operations, which may have economic implications in terms of grafting percentage and fruit yield. The response of Solanaceous plants to graft incompatibility may differ based on the combination of the scion and the rootstock selected. Severe incompatibilities have been observed in, for example, tomato/pepper (scion/rootstock) grafts, while moderate incompatibilities have been observed in eggplant and tomato (scion/rootstock) grafts. This is related to yield and the number of grafted plants that survived after grafting [32].
Rootstock regrowth, also referred to as “suckering” or adventitious bud growth, usually occurs about 14 days after grafting success. The regrowth becomes vigorous and occurs beneath the graft union on the rootstock. Usually both rootstocks (S. macrocarpon and S. aethiopicum) exhibit adventitious bud regrowth (Figure 2).
Grafted tomato plants showing adventitious bud regrowth (red arrow). Picture by Charles Agyeman.
Monocotyledonous plants cannot be grafted because they lack the ability to form cambium layers, compared to dicotyledonous plants. Temperature and relative humidity levels are crucial environmental factors for graft union formation, and acclimatization of grafted plants. The regulation of these post-grafting factors will influence the survival rates of the grafted plants, grafting success, and yield. Generally, a higher relative humidity in the grafting chamber tends to favor grafted tomato plants, as grafted plants do not lose moisture at higher rates [33]. High humidity within the grafting chamber can be achieved by misting the chamber regularly with water; the use of plastic polythene to cover the grafting chamber acts as an insulator, which shields the plants from the changes in temperature and other weather conditions.
An ideal post-grafting operation should therefore include the maintenance of an ideal air temperature and relative humidity of 25–28°C and 80–90%, respectively, which will promote a higher survival rate and quality of grafted seedlings [34]. In situations where temperature levels have exceeded 30–32°C, the leaf weights (dry weight and fresh weight) have been reported to reduce significantly in watermelon [35].
Quality has become the hallmark of consumers who purchase vegetables as part of their daily dietary requirements; consumers therefore use certain visual and nonvisual attributes to determine the quality of vegetables and fruits in general. Consumers determine the quality of tomato fruits based on their appearance (size, color, and shape) and texture (firmness, mealiness, and juiciness) as well as their flavor and nutritional content [36]. However, different market players along the vegetable value chain their standard for quality. The quality of tomato is based on soluble solids, acidity, sugars, pH, and shelf life [37].
Vegetable farmers and traders prefer tomato cultivars which exhibit firmness and can withstand mechanical damage, whilst in transit to various market centers [38]. The term fruit quality, which can be defined based on the visual and sensory properties such as color and sweetness, has been found to be controlled by certain inherent genes in some plant cultivars; some of these genes or genetic traits can be bred into new genotypes from other wild species [39].
Conflicting reports on the influence of grafting on fruit quality in vegetables exist. Positive and negative influences of grafting have been documented [40]. In their review of the impact of grafting on fruit quality in vegetables, Rouphael et al. [40] attributed these conflicting results to the differences in environments, production methods, scion/rootstock combinations, and harvest dates.
In an experiment conducted by Matsuzoe et al. [41], where tomatoes (Momotaro) were grafted on three Solanum species (S. torvum, S. toxicarum and S. sisymbriifolium), there were, however, no significant differences in the quality of grafted and ungrafted tomatoes in relation to the amount of sugars and their organic acid contents.
Traditionally, field and pot screening have been used to identify plant cultivars that are resistant to root-knot-nematodes as screening of rootstocks against root-knot nematodes is essential for every grafting program, because this informs the selection of the right rootstock for grafting. In a field experiment to evaluate the performance of grafted eggplant cultivars on wild Solanum rootstocks against root-knot nematodes, results revealed that the wild Solanum rootstocks S. torvum, S. sisymbriifolium, and S. khasianum were resistant to root-knot nematode when inoculated with 1000 nematode juveniles [42]. The non-grafted plants generally flowered before the grafted plants, a situation which is attributed to the cut back of the leaves of the scion to reduce transpiration which slowed down the rate of growth. Thirty-three tomato genotypes screened for root-knot nematode resistance under five inoculum levels (100, 500, 1000, 1500, and 2000) showed increasing inoculum level with corresponding increase in gall score and fresh root weight [43]. Among the 33 tomato genotypes tested, Mongal F1 T-11 had the lowest mean gall score of 3.25 and “Beef Master” had a value of 3.75 with reproductive factors of 0.71 and 0.53, respectively. Tomato cultivars that are resistant to root-knot nematodes have a reproductive factor less than one, which implies that the plant is able to suppress the reproduction cycle of the organism once it gains entry into the roots [44]. In a grafting study by Agyeman [19], significant differences were not observed among total soluble solids (TSS), pH, and titrable acidity (TA) for the tomato variety “Pectomech” grafted onto S. aethiopicum and S. macrocarpon after infection to 500 and 1000 nematodes per pot (Table 3).
Treatments | Inoculum levels | TSS | TSS/TA | pH | TA |
---|---|---|---|---|---|
P/SA | 0 | 5.42 | 3.04 | 4.47 | 1.87 |
P/SM | 0 | 5.99 | 4.02 | 4.36 | 1.65 |
P/SA | 500 | 6.27 | 4.01 | 4.55 | 1.67 |
P/SM | 500 | 6.33 | 4.40 | 4.79 | 1.62 |
P/SA | 1000 | 6.67 | 4.27 | 4.42 | 1.65 |
P/SM | 1000 | 5.78 | 6.44 | 4.72 | 1.22 |
P/SA | 5000 | 6.3 | 4.25 | 4.45 | 1.45 |
P/SM | 5000 | 6.28 | 3.81 | 4.43 | 1.66 |
LSD(P = 0.05) | ns | ns | ns | ns |
Comparison of grafted rootstocks and inoculum level interaction on TSS, TSS/TA, pH, and TA.
P/SA = Pectomech grafted onto Solanum aethiopicum; P/SM = Pectomech grafted onto Solanum macrocarpon; TSS = Total soluble solids; TA = Titrable acidity; LSD = Least significant difference; ns = no significant difference. Agyeman [19].
In a pot culture experiment conducted by Dhivya et al. [45], 10 Solanum plant genotypes (S. torvum, S. incanum, S. xanthocarpum, S. aethiopicum, S. sisymbrifolium, S. viarum, S. violaceum, Physalis peruviana, and TNAU Tomato Hybrid CO-3 and US-618) consisting of eight wild species and two F1 cultivars were evaluated for their resistance to root-knot nematode over a 60-day period, and the results showed that S. sisymbrifolium rootstock had the highest shoot fresh weight and dry weight of 103.87 and 10.44 g, respectively.
The rootstocks, S. sisymbrifolium, Physalis peruviana, and S. torvum recorded the least nematode population of 39, 40 and 43 per 200 cc of soil and a reproductive factor of 0.71, 0.74, and 0.84, respectively. Solanum sisymbrifolium, P. peruviana, and S. torvum were resistant to root-knot nematode (Meloidogyne incognita), and S. incanum and S. aethiopicum were found to be moderately resistant to Meloidogyne incognita.
The resistance offered by plants to the damage caused by root-knot nematodes have been well researched and attributed to the presence of a single dominant gene (Mi gene). The Mi gene confers resistance to various root-knot nematode species (M. incognita, M. javanica, and M. arenaria) in addition to whiteflies and aphids [46]. Solanum spp., for example, Lycopersicon peruvianum and S. torvum have been reported to have this resistant gene, which enables the plant to tolerate the feeding activities and the reproductive abilities of root-knot nematodes [47]. The Mi gene was first discovered in an accession of a wild L. peruvianum in South America from which commercial F1 varieties were introgressed with the gene [48]. This process involves the extraction and detection of the gene using DNA markers and subsequent isolation of the gene for introgression. In other related research conducted using the positional cloning approach to isolate gene with linked traits and the subsequent sequencing of the DNA, Kaloshian et al. [49] reported that the sequencing analysis showed two genes, which were identical to each other (Mi-1.1 and Mi-1.2), which also confers resistance to three species of root-knot nematodes namely M. arenaria, M. javanica, and M. incognita.
Several DNA markers have been developed for the detection of the Mi gene in plants using polymerase chain reaction (PCR) amplifications. Devran et al. [50] screened for the Mi gene using gene specific primers C1/2 (5′-cagtgaagtggaagtgatga-3′) and C2S4 (5′-ctaagaggaatctcatcacagg-3′) for screening F2 tomato plants for the root-knot nematode resistance gene. A 1.6 kb amplification product was amplified in these containing the Mi-1.2 gene in the 3′ region; however, it was found to be absent in the susceptible F2 plants.
Similarly, in another study, the Mi-1.2 gene was introgressed into S. melongena to confer resistance to M. javanica and aphids. The study revealed that the transgenic eggplant was able to confer resistance to M. javanica but not aphids [51]. In confirming the presence of the Mi-1.2 gene in the transgenic eggplant, a reverse-transcription polymerase chain reaction assay with the Mi specific primers C2D1 (5′-ctagaa agtctgtttgtgtctaacaaagg-3′) and C2S4 (5′-ctaagaggaatctcatcacagg-3′) amplified a single PCR band of 915 bp, which was present but absent in the nontransgenic S. melongena.
A study in Morocco by Mehrach et al. [52] to detect the Mi-1.2 gene in 14 begomovirus-resistant breeding lines with known resistance was also undertaken using a two-step PCR approach. The primer pairs PM3Fb/PM3Rb and REX primers used in a multiplex PCR amplified a band of 720 bp for both susceptible and resistant varieties; however, the resistant varieties (Motelle and Better Boy) showed an additional band of 500 bp, indicating the presence of the Mi gene in those cultivars.
In distinguishing between heterozygous and homozygous plant cultivars with the Mi-1.2 gene, the primer pairs of PMiF3/PMiR3 amplified a single unique band of 350 bp for the susceptible cultivars (Moneymaker and Daniella). However, 550 and 350 bp fragments for both the homozygous and heterozygous plant resistant cultivars “Motelle” and “Better Boy” were amplified, respectively.
Farmers are the ultimate beneficiaries of grafted plants; therefore, healthy grafted seedlings production is important at affordable prices. The high costs involved in the grafting process are due to high labor requirements, grafting input costs, and seeds of rootstock. These associated costs therefore limit the usage of grafted plants by growers or farmers. Grafting costs can be reduced through training of selected farmers from farmer groups, who will in turn train other farmers (trainer of trainers). Information related to this technology can be passed on to farmers and other interested stakeholders through extension programs, for example, workshops, fairs, field days, and on-farm trials. There is also the need for undertaking extensive disease diagnosis in specific areas and feedback given to farmers. Tomato rootstock breeding efforts can lead to production of rootstocks to specific environments, pests and diseases, and other abiotic stresses.
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\n\nOur reputation – Everything we publish goes through a two-stage peer review process. We’re proud to count Nobel laureates among our esteemed authors. We meet European Commission standards for funding, and the research we’ve published has been funded by the Bill and Melinda Gates Foundation and the Wellcome Trust, among others. IntechOpen is a member of all relevant trade associations (including the STM Association and the Association of Learned and Professional Society Publishers) and has a selection of books indexed in Web of Science's Book Citation Index.
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