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Tiefenbacher",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/10765.jpg",keywords:"Managing Urbanization, Managing Development, Managing Resource Use, Drought Management, Flood Management, Water Quality Monitoring, Air Quality Monitoring, Ecological Monitoring, Modeling Extreme Natural Events, Ecological Restoration, Restoring Environmental Flows, Environmental Management Perspectives",numberOfDownloads:24,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"January 12th 2021",dateEndSecondStepPublish:"February 9th 2021",dateEndThirdStepPublish:"April 10th 2021",dateEndFourthStepPublish:"June 29th 2021",dateEndFifthStepPublish:"August 28th 2021",remainingDaysToSecondStep:"2 months",secondStepPassed:!0,currentStepOfPublishingProcess:4,editedByType:null,kuFlag:!1,biosketch:"A geospatial scholar working at the interface of natural and human systems, collaborating internationally on innovative studies about hazards and environmental challenges. Dr. Tiefenbacher has published more than 200 papers on a diverse array of topics that examine perception and behaviors with regards to the application of pesticides, releases of toxic chemicals, environments of the U.S.-Mexico borderlands, wildlife hazards, and the geography of wine.",coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"73876",title:"Dr.",name:"John P.",middleName:null,surname:"Tiefenbacher",slug:"john-p.-tiefenbacher",fullName:"John P. Tiefenbacher",profilePictureURL:"https://mts.intechopen.com/storage/users/73876/images/system/73876.jfif",biography:"Dr. John P. Tiefenbacher (Ph.D., Rutgers, 1992) is a professor of Geography at Texas State University. His research has focused on various aspects of hazards and environmental management. 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1. Introduction
Various types of lubricants are available all over the world including mineral oils, synthetic oils, re-refined oils, and vegetable oils. Most of the lubricants which are available in the market are based on mineral oil derived from petroleum oil which are not adaptable with the environment because of its toxicity and non-biodegradability [1, 2]. Unknown petroleum reserve and the increasing consumption, which made concern to use petroleum based lubricant thus, to find the alternative lubricant to meet the future demand is an important issue [3]. Therefore, vegetable oil can be played a vital role to substitute the petroleum lubricant as it possesses numerous advantage over base lubricant like renewability, environmentally friendly, biodegradability, less toxicity and so on [4-8]. It has been reported that yearly 12 million tons of lubricants waste are released to the environment [9]. However, it is very difficult to dispose it safely for the mineral oil based lubricants due its toxic and non-biodegradable nature. To reduce the dependency on petroleum fuel, legislations have been passed to use certain percentage of biofuel in many countries, such initiative also required for lubricant as well [10]. Vegetable oils are mainly triglycerides which contain three hydroxyl groups and long chain unsaturated free fatty acids attached at the hydroxyl group by ester linkages acids favors triglycerides crystallization [11, 12]. The unsaturated free fatty acid which is defined as the ratio and position of carbon-carbon double bond, one two and three double bonds of carbon chain is named as a oleic, linoleic, and linolenic fatty acid components respectively [13]. The main limitations of vegetable oil are its poor low temperature behavior, oxidation and thermal stability and gumming effect [14, 15]. These stabilities and pour point behavior can be ameliorated by transesterification. Moreover the inferior flow property does not affect much in the tropical countries. Quinchia et al. [16] stated that, improving the potentiality of biolubricants some technical properties including available range of viscosities are need to improved. To do so, environmentally friendly viscosity modifier can be used. viscosity is the most important property for the lubricants since it determines the amount of friction that will be encountered between sliding surfaces and whether a thick enough film can be build up to avoid wear from solid-to-solid contact. Since little chance of viscosity with fluctuations in temperature is desirable to keep variations in friction at a minimum, fluid often are rated in terms of viscosity index. The less the viscosity is changed by temperature, the higher the viscosity index. Ethylene–vinyl acetate (EVA) and styrene–butadiene–styrene (SBS) copolymers were used to increase the viscosity range of high-oleic sunflower oil, in order to design new environmentally friendly lubricant formulations with increased viscosities. The maximum kinematic viscosities, at 40 and 100 ◦C, were increased up to around 150–250 cSt and 26–36 cSt, respectively [17].
Despite of having lot of advantages of biolubricant over petroleum based lubricant, the attempt to formulate the biolubricant and its applications are very few. Thus, in this article we sought to extend our investigation and to test the tribological characteristics and compatibility of non-edible Jatropha oil based biolubricant for the automotive application. The reason of selecting Jatropha oil as a base stock is it does not contend with the food and can be grown in marginal land.
2. Experimental
2.1. Lubricant sample preparation
There were six different types of lubricant sample were investigated in this study. The lubricant SAE 40 was used as a base lubricant and comparison purpose. Others samples were prepared by mixing of 10%, 20%, 30%, 40% and 50% Jatropha oil in SAE 40. The samples were mixed with the base lubricant by a homogeneous mixture machine.
2.2. Friction and wear evaluation
The apparatus used in the friction and wear testing process were Cygnus Friction and Wear Testing Machine which is connected with a personal computer (PC) with data acquisition system. It is a tri-pin-on-disc machine which is conducted by using three pins on a disc as testing specimens. Specifications of the Cygnus Test Machine are tabulated in Table 1. The block diagram of friction and wear testing are shown in Fig. 1. During the test the load of 30N and rotational speed of 2000 rpm were applied on pin.
Parameter
Value
Test Disc Diameter
110.0 mm
Test Pin Diameter
6.0 mm
Test Disc Speed Range
25 to 3000 rpm
Motor
Tuscan; (2000 rpm, 1.5 kW)
Load Range
0 KG to 30 KG
Electrical Input
220 Volt AC 50 Hz
Table 1.
Specification of Cygnus wear testing machine
Figure 1.
Block Diagrams of Friction and Wear Testing
2.3. Preparation of the specimen
The specimens were prepared from aluminum and cast iron material. Aluminum was used to build three pin and cast iron is used for disc specimen. The construction geometry and the dimension are shown in Fig. 2. Prior to conduct the test it was ensured that the surface of the specimens are cleaned properly i. e, free from dirt and debris. Alcohol was used for cleaning purpose.
Figure 2.
Schematic diagram of the experimental set up and dimensions geometry’s of pins and disc specimen
2.4. Lubricant analyses
Multi element oil analyzer (MOA) was used to measure the wear elements in the lubricants by Atomic Emission Spectroscopy (AES). Whereas, for viscosity measurement the automatic Anton Paar viscosity meter was used with standard ASTM D 445. Viscosity was measured for both 40°C and 100°C controlled bath temperatures.
3. Results and discussion
3.1. Friction and wear characterization
Fig. 3 show the pins wear as a function of sliding time for various Jatropha oil blended biolubricants. At the operating condition of 2000 rpm and 30 N loads, the linear pin wear varied from 0.02 to 0.05 mm. It is observed that the maximum wear occurred in the beginning of the experiment using biolubricants. It can be seen form the Fig. 3, that the maximum wear was occurred for JBL40 while the minimum wear was observed for JBL10. The results can be attributed to the maximum ability of the JBL 10 biolubricant film to protect metal to metal contact and keep consistency throughout the operation time while this ability is least for JBL40. It can also be seen that the rate of wear throughout the time is almost identical for the biolubricants whereas, the reducing trend is observed for the base lubricant. At the beginning of the test, the wear rate was very fast for few minutes which are known running-in period. During this period, the asperities of the sliding surface are cut off and the contact area of the sliding surface grows to an equilibrium size. After certain period of time, equilibrium wear condition between pins and disc surface was established and thereby the wear rate became steady. It can be identified from the Fig. 3 that the biolubricants JBL 30, JBL 40 and JBL 50 showed high wear while base lubricant, JBL 10 and JBL 20 impart low pin wear and their values are nearly same with each other.
Fig. 4 sows the loos of material from the pin for different percentage of biolubricant samples. It seems quite clear that the loos of material from the pins are highest for 50% biolubricant and that is least for base lubricant. It can also be interpreted that the loos of material from JBL 10 is almost similar with base lubricant and this loos of material is increasing with increasing biolubricant percentages.
Figure 3.
The linear pin wear as a function of sliding time for various Jatropha oil biolubricants.
Figure 4.
Loos of material form the pin for various biolubricant percentages
3.2. Coefficient of friction
Fig.5 shows the friction coefficient plotted against the sliding time for various Jatropha oil biolubricants. The results of the figure depict that the lubricant regime that occurred during the experiment were the boundary lubrication with the value of friction coefficient for boundary lubricant in the range of 0.001 to 0.2 except for 50% of Jatropha oil biolubricant. For JBL 0, it can be seen that the coefficient of friction is highest at the beginning and then it fell down rapidly and became least with compared to all tested samples after half of the operation time. The biolubricant percentage from 10 to 40% showed likely to be similar coefficient of friction (µ) which is almost 0.15. Whereas, the 50 % added Jatropha oil showed the coefficient of friction value of ~ 0.225 throughout the operation time. The fatty acid component of biolubricants formed multi and mono layer on the surface of the rubbing zone and make stable film to prevent the contact between the surfaces.
Figure 5.
The Coefficient of friction as a function of sliding time for various Jatropha oil biolubricants
3.3. Lubricants temperature
Fig.6 shows the relationship of the averages oil temperature of varies percentage of Jatropha oil biolubricants with the sliding time. The rise of temperature during the running hour (1 h) for JBL 10 is least while the highest change is occurred for JBL 40 which is 11.77°c and 25.49°C respectively. The temperature rises of other samples are of 12.8°C, 18.65°C and 13. 66°C for 20% 30% and 50% Jatropha oil added biolubricants respectively. The results of the Fig. 6 show that the JBL 10 has the highest potentiality to retain its property without much changing its temperature. From the figure it can also be interpreted that up to 30 minutes rate of change of temperature is high while the changing rate is low for second half of the operation time. It can be explained that during second half of the operation time heat produced in the lubricant due friction and the heat dissipated to the outside is nearly equilibrium.
Figure 6.
The Lubricant Temperature as a function of sliding time for various Jatropha oil biolubricants
3.4. Viscosity
Viscosity is the measure of resistance to flow [18]. Table 2 shows the viscosity grade requirement for the lubricants set by International standard organization (ISO), while Fig. 7 shows the viscosity of tested different biolubricant samples. The comparison of the results of the Fig.7 with that of ISO grade illustrates that in case of 40°C, the biolubricants JBL 40 and JBL 50 did not meet the ISO VG100 requirement. On the other hand all other biolubricants meet the entire ISO grade requirement as well. It can also be noted that the viscosity of biolubricants are much higher than standard requirements
The viscosity of various percentages of biolubricants at 40°C and 100°C
3.5. Elemental analysis
The aim of the elemental analysis by using Multi Element Oil Analyzer (MOA) is to determine the kinds and amount of metal contain in the lubricating oil. Table 3 shows the elemental analysis of tested lubricant sample by using MOA before and after the test. From the Table 3, it can be noticed that the base lubricant contains higher Silver (Ag), Zinc (Zn), Phosphorus (P), Magnesium (Mg) and Boron (B) with in high percentage compared to other element while, in pure Jatropha oil, Calcium (Ca) and Silicon (Si) are the higher element compared with other element. Some of the elements are used as additive in the lubricant to ameliorate the lubricants tribological properties. From the results, increasing number of iron (Fe) and aluminum (Al) molecules are observed with increasing percentages of Jatropha oil in the base lubricants. The source of Fe and Al are mainly cast iron plate and aluminum plate. Due to lower hardness of the aluminum pin the extraction of aluminum molecule form the pin is much higher than cast iron plate. The changes of other elements were observed before and after the test. It is clear from the elemental analysis that, most of elements were decreased after the test, by oxidizing and the chemical interaction among the elements.
Table 3.
Elemental analysis of tested lubricant sample
3.6. Surface texture analysis
There are various types of wear in the mechanical system, such that abrasive wear, adhesive wear, fatigue wear and corrosive wear. Since the lubricant regime occurred in this experiment was boundary lubrication thereby, abrasive wear, adhesive wear, fatigue wear and corrosive wear were observed in to the rubbing zone. All these wears mechanisms found in this experiments but the mostly the wear phenomenon were abrasive and adhesive wear. This is because of an existence of straight grooves in the direction of the sliding direction. These grooves exist because the asperities on the hard surface (disc) touched the soft surface (pins) and hade a close relationship with the thickness of lubrication film. The optical images of the tested cast iron plate using various types of biolubricants are shown in Fig. 8. Referring to the Fig. 8, it is found that the wear increases with increasing percentage of Jatropha oil in the biolubricants. Reduction of lubricant film thickness leads to the surfaces to come closer to each other and cause higher wear.
Figure 8.
Optical image of the surface of the cast iron plate for different biolubricants (magnification 30 ×): (a): JBL 0, (b): JBL10, (c): JBL 20, (d): JBL 30, (e): JBL 40, (f): JBL 50
4. Conclusions
Based on the experimental study the following conclusion can be drawn:
The rates of wear for various percentage of biolubricant were different. Moreover the wear rate for 10% Jatropha added biolubricants were almost identical with base lubricant.
Lower the resistance to wear, higher coefficient of friction.
At the beginning of the test rate of wear as well as rise in temperature were high. With respect to wear rate and rise in temperature during entire operation time, the JBL 10 biolubricant showed best performance in terms of its ability to withstand its properties.
From the elemental analysis of the biolubricants, it is found, Iron and Aluminum were increased after the test due to the loos of material from the pin and the disc while, some element like Phosphorus, Calcium and Magnesium were decreased by oxidizing and due to other chemical interaction.
In terms of viscosity, almost all biolubricants met the ISO viscosity grade requirement whereas, 40% and 50% addition of Jatropha oil do not meet the ISO VG 100 requirement at 40°C.
According to the experimental result, it can be recommended that the addition of 10% Jatropha oil in the base lubricant is the optimum for the automotive application as it showed best overall performance in terms of wear, coefficient of friction, viscosity, rise in temperature etc.
Acknowledgement
The authors would like to acknowledge the Department of Mechanical Engineering, University of Malaya, Ministry of Higher Education (MOHE) of Malaysia for HIR grant (Grant No. UM.C/HIR/MOHE/ENG/07) and ERGS grant no ER022-2011A which made this study possible.
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Masjuki",slug:"h.h.-masjuki",email:"masjuki@um.edu.my",position:null,institution:{name:"University of Malaya",institutionURL:null,country:{name:"Malaysia"}}},{id:"152617",title:"M.Sc.",name:"Md. Shahauddin",middleName:null,surname:"Ahmmad",fullName:"Md. Shahauddin Ahmmad",slug:"md.-shahauddin-ahmmad",email:"shahabuddin.suzan@yahoo.com",position:null,institution:null},{id:"152618",title:"Mr.",name:"Md.Mofijur",middleName:null,surname:"Rahman",fullName:"Md.Mofijur Rahman",slug:"md.mofijur-rahman",email:"mofijduetme@yahoo.com",position:null,institution:null},{id:"166960",title:"Dr.",name:"N.W.M.",middleName:null,surname:"Zulkifli",fullName:"N.W.M. Zulkifli",slug:"n.w.m.-zulkifli",email:"nurinmz@um.edu.my",position:null,institution:null}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Experimental",level:"1"},{id:"sec_2_2",title:"2.1. Lubricant sample preparation",level:"2"},{id:"sec_3_2",title:"2.2. Friction and wear evaluation",level:"2"},{id:"sec_4_2",title:"2.3. Preparation of the specimen",level:"2"},{id:"sec_5_2",title:"2.4. Lubricant analyses",level:"2"},{id:"sec_7",title:"3. Results and discussion",level:"1"},{id:"sec_7_2",title:"3.1. Friction and wear characterization",level:"2"},{id:"sec_8_2",title:"3.2. Coefficient of friction",level:"2"},{id:"sec_9_2",title:"3.3. Lubricants temperature",level:"2"},{id:"sec_10_2",title:"3.4. Viscosity",level:"2"},{id:"sec_11_2",title:"3.5. Elemental analysis",level:"2"},{id:"sec_12_2",title:"3.6. Surface texture analysis",level:"2"},{id:"sec_14",title:"4. Conclusions",level:"1"},{id:"sec_15",title:"Acknowledgement",level:"1"}],chapterReferences:[{id:"B1",body:'SalihN.SalimonJ.YousifE.Synthetic biolubricant basestocks based on environmentally friendly raw materials. Journal of King Saud University-Science 2011'},{id:"B2",body:'AdhvaryuA.LiuZ.ErhanS.Synthesis of novel alkoxylated triacylglycerols and their lubricant base oil properties. Industrial Crops and Products 200521113119'},{id:"B3",body:'ShahabuddinM.MasjukiH. H.MAKalamet.alEffect of Additive on Performance of C.I. Engine Fuelled with Bio Diesel. Energy Procedia 20121416241629'},{id:"B4",body:'SiniawskiM. T.SanieiN.AdhikariB.DoezemaL. A.Influence of fatty acid composition on the tribological performance of two vegetable-based lubricants. Journal of Synthetic Lubrication 200724101110'},{id:"B5",body:'Salunkhe DK. World oilseeds: chemistry, technology, andutilization. 1992'},{id:"B6",body:'Hwang HS, Erhan SZ.Lubricant base stocks from modified soybean oil. AOCS Press: Champaign, IL; 2002'},{id:"B7",body:'IngT. C.RafiqA. K. M.SyahrullailS.Friction Characteristic of Jatropha Oil using Fourball Tribotester. In: Regional Tribology Conference- RTC2011. Langkawi, Kedah, Malaysia: 2011'},{id:"B8",body:'ShahabuddinM.KalamM. A.MasjukiH. H.MofijurM.Tribological characteristics of amine phosphate and octylated/butylated diphenylamine additives infused biolubricant. Energy Education Science and Technology Part A: Energy Science and Research 20123089102'},{id:"B9",body:'TottenG. E.WestbrookS. R.ShahR. J.Fuels and Lubricants Handbook: Technology,Properties, Performance, and Testing. 2003885909p.'},{id:"B10",body:'Liaquat AM, Masjuki HH, Kalam MA et al.Application of blend fuels in a diesel engine. Energy Procedia 20121411241133'},{id:"B11",body:'JayadasN.NairK. P.Coconut oil as base oil for industrial lubricants--evaluation and modification of thermal, oxidative and low temperature properties. Tribology international 200639873878'},{id:"B12",body:'FoxN.StachowiakG.Vegetable oil-based lubricants-a review of oxidation. Tribology international 20074010351046'},{id:"B13",body:'WaleskaC.DavidE. W.KraipatC.JosephM. P.The effect of chemical structure of base fluids on antiwear effectiveness of additives. Tribol. Int. 2005383216'},{id:"B14",body:'PonnekantiN.KaulS.Development of ecofriendly/biodegradable lubricants: An overview. 2012'},{id:"B15",body:'MofijurM.MasjukiH. H.MAKalamet.alPalm Oil Methyl Ester and Its Emulsions Effect on Lubricant Performance and Engine Components Wear. Energy Procedia 20121417481753'},{id:"B16",body:'QuinchiaL.DelgadoM.ValenciaC.et al.Viscosity modification of different vegetable oils with EVA copolymer for lubricant applications. Industrial Crops and Products 201032607612'},{id:"B17",body:'QuinchiaL.DelgadoM.ValenciaC.et al.Viscosity modification of high-oleic sunflower oil with polymeric additives for the design of new biolubricant formulations. Environmental science & technology 20094320602065'},{id:"B18",body:'ShahabuddinM.MAKalamMasjuki. H. H.et al.An experimental investigation into biodiesel stability by means of oxidation and property determination. Energy 2012'},{id:"B19",body:'Rudnick LR. Automotives Gear Lubricants, Synthetics, mineral oils, and bio-based lubricants: chemistry and technology. Taylor and Francis,Florida; 2006'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"M. Shahabuddin",address:null,affiliation:'
Centre for Energy Sciences, Faculty of Engineering, University of Malaya, Kuala Lumpur, Malaysia
Centre for Energy Sciences, Faculty of Engineering, University of Malaya, Kuala Lumpur, Malaysia
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1. Introduction
A fundamental biotechnological method, genetic engineering, enables the production of plants with desired properties. Further breeding of obtained plants is possible by another biotechnological tool of great importance such as in vitro culture techniques. Applying of tissue culture procedures enables regeneration and multiplication of desired plants in controlled conditions in short time periods. Developing of plant tissue cultures also allows preservation and improvement of plant species endangered in nature, especially medicinal plant species such as e.g. common centaury (Centaurium erythraea Rafn.).
1.1 Genetic transformations of plants
Genetic engineering of plants represents a unique field of biotechnology where genes and other DNA sequences are manipulated to obtain plants with certain desired properties. The necessary steps in genetic engineering are the isolation and modification of the gene of interest, the vector construction, transformation and the final selection of transgenic plants. Genetic transformation of plants implies stable incorporation of foreign genes into the plant genome [1]. Genetic transformation of plants can be very successfully achieved in two ways:
Techniques of direct gene transfer (microbombardment - biolistic method, microinjection, electroporation, polyethylene glycol mediated transformation, protoplast fusion)
Gene transfer techniques through biological vectors (viruses or bacteria - Agrobacterium rhizogenes and Agrobacterium tumefaciens)
Direct gene transfer techniques are continually developing and improving over the years. However, a considerable variation in the stability, integration and expression of the introduced gene represent their significant deficiencies. On the other hand, techniques of gene transfer through bacteria of the genus Agrobacterium have proven to be very efficient and straightforward. These techniques allow a more precise and stable integration of the desired gene into the plant genome [2].
1.1.1 Plasmids of bacteria of the genus Agrobacterium
The Agrobacterium genus is a group of Gram-negative soil bacteria. Unlike the wild type, laboratory Agrobacterium strains are commonly used for genetic transformations of numerous plant species [3]. Different plant species show different susceptibility to infection by Agrobacterium bacteria [4]. In general, dicotyledonous plants are more susceptible to infection by Agrobacterium bacteria than monocotyledons. Bacteria of the genus Agrobacterium can be present in the vascular tissue of plants (xylem and phloem) but without showing disease symptoms. It indicates that the interaction of bacteria and plant cells does not always occur. Infection and pathological changes of plant tissue occuronly after injury of the plant cells when released phenolic compounds causea positive chemotoxic reaction in Agrobacterium. These phenolic compounds further induce genes, located on chromosomal and plasmid DNA, responsible for bacterial virulence. During genetic transformations, the plasmids in bacterial cells serve as vectors. Bacteria of the genus Agrobacterium have large conjugated plasmids (200–250 kb), parts of which have been successfully integrated into the plant genome during transformation processes for almost half a century [5]. In these bacteria, two basic types of plasmids are identified by the kind of disease they cause: Ti-plasmid (tumor-inducing) in A. tumefaciens and Ri-plasmid (root-inducing) in A. rhizogenes.
Both types of plasmids contain three genetic regions that are necessary for the genetic transformation process:
T-DNA (transferred DNA) – a part of plasmid DNA, size 10–30 Kbp. The transport of T-DNA into a plant cell allows expression of bacterium genes in the plant itself [6].
Vir (virulence) region, size 35 Kbp, which consists of several large loci (virA, virB, virC, virD, virE, virG, virH and virJ). Vir genes encode the synthesis of vir proteins, which are responsible for the process of recognizing specific compounds synthesized by a plant cell after injury and transferring T-DNA to the host cell.
The bacterial chromosome region, which is made up of chv (chromosomal virulence) genes that regulate bacterial chemotaxis and bacterial binding to an injured plant cell [7]. According to Simonović [8], the chvA and chvB genes are responsible for the production and secretion of cyclic 1,2-glycans while chvE product forms a transmembrane receptor complex together with the plasmid virA product.
The development of plant tumor tissue occurs after a part of the plasmid DNA (T-DNA) is transported and incorporated into the DNA of the plant. T-DNA of Ti- and Ri-plasmids carries oncogenes, the genes responsible for tumor formation and encoding enzymes that catalyze the synthesis of auxins and cytokinins. Transcription of these genes leads to an increased amount of these plant hormones, which stimulates uncontrolled cell division, tumor development and mass proliferation of transformed cells [9].
1.1.2 Mechanism of genetic transformation of a plant cell using Agrobacterium tumefaciens
Successful genetic transformations of plants using Agrobacterium require a bacterial strain with appropriate density, explant tissue culture, determination of the substrate and time for cocultivation, as well as temperature and selection regime [10]. In wild strains of Agrobacterium, the Ti- and Ri- plasmids have a T-DNA region divided into two sequences (TL and TR) between which is a TC sequence that is not transmitted to the plant genome [11]. Oncogenes are located on the TL sequence while genes for opine synthesis are located on the TR sequence. Opines accumulate in transformed plant cells thus representing metabolic markers for genetic transformations. They seem useless for the plant as no appropriate enzymes for their degradation have been found in plant tissues. Pathogenic bacteria use released opines as a source of carbon and nitrogen. The genes on the plasmid itself encode the enzymes for catabolism of opines whereby the opine, whose synthesis is induced in transformed tissue, is degraded [12]. In laboratory strains of Agrobacterium, natural T-DNA can be replaced by a gene of interest that should be introduced into the plant genome by genetic transformation.
Genetic transformation of plants by Agrobacterium is a process that, in general, consists of three steps:
Recognition and “attack” of a plant cell by a bacterium
Injured plant cells begin to synthesize specific compounds (amino acids, sugars and organic acids). In dicotyledons these are phenolic derivatives syringone and acetosyringone [13] while in monocotyledons it is ethyl ferrulate [14]. These compounds cause a positive chemotoxic reaction that leads to interaction of plant cells and bacteria. Binding of a bacterium to the plant cell represents the first step in the mechanism of genetic transformation.
Transfer of T-DNA into a plant cell
VirD and VirE proteins play a significant role in the formation of single-stranded T-DNA, its cutting and transfer to the plant cell. The products of the VirD region (VirD1 and VirD2 proteins) are part of an endonuclease complex catalyzing specific single-stranded breaks that release a single-stranded T-DNA segment from the Ti plasmid. After excision, the T-chain shifts polarly and covalently binds the VirD2 protein. This protein protects the T-chain from destruction and remains there during transport to the plant cell. The transport of VirD2/T-DNK complex (immature T-complex) into the cytoplasm of the host cell, together with several other vir proteins (VirE2, VirF and VirE3), is performed through a specialized membrane transport system. In cytoplasm of the plant cell, a mature T-complex is formed, which is protected from the action of nucleases during transport to the nucleus [15].
T-DNA gene incorporation and expression
The last, and probably the most critical step of genetic transformation, is the integration of T-DNA into the host genome. Integration into the plant genome occurs by non-homologous recombination in any part of the genome, taking preferentially place in regions with a higher percentage of adenine-thymine (AT) bonds [16]. Transgene expression depends on numerous factors such as e.g. the number of copies of an introduced gene and the site of transgene integration [17, 18, 19]. Stably integrated T-DNA is transferred to the next generation [20].
The mechanism of genetic transformation today involves the use of so-called “disarmed” A. tumefaciens strains, in which oncogenes are removed from the plasmids. The system of binary vectors usually used in genetic transformation processes divides the original Ti-plasmid into two plasmid vectors: an artificial (recombinant) Ti-plasmid carrying the gene of interest and an auxiliary plasmid [21]. In each binary plasmid, additionally to a foreign gene of interest the marker genes are also inserted and divided into selective and reporter genes [22]. The selective marker genes are most often prokaryotic genes that encode resistance to some antibiotics (kanamycin, hygromycin etc.). This negative selection implies using a selective medium which allows the distinction of transformed plant cells from non-transformed ones. Reporter genes are common but not necessary parts of the binary vector. These genes encode products which can be involved in the monitoring of transformation process (growth of transformed cells, regeneration efficiency and growth of transgenic plants). Visual reporter genes have been increasingly used because they do not require special artificial substrates for detection, and their expression is easily visually detected in tissue without destroying.
1.2 Cytokinins
Cytokinins (CKs) are plant hormones that individually or in combination with other hormones regulate numerous developmental and physiological processes in plants [23]. The name cytokinins comes from their originally discovered function, which is the stimulation of cell division – cytokinesis [24]. Cytokinins are divided into two major groups: natural and synthetic.
Natural CKs by chemical structure represent adenine derivatives. Depending on the component that binds to the adenine ring, natural CKs are further categorized into two groups:
Isoprenoid CKs – appear when one isoprenoid C5 unit binds to the N6 atom of adenine [25]. Natural isoprenoid CKs include N6-(∆2-isopentenyl)adenine (iP), trans-zeatin (tZ), cis-zeatin (cZ) and dihydrozeatin (DHZ) and their derivatives (ribosides, N- and O-glucosides, ribotides) differing in their biological functions. The chemical structures of isoprenoid cytokinins are shown in Figure 1. The contents and distribution of individual isoprenoid CKs in plants vary depending on the plant species, particular plant tissue, developmental stage, etc.
Aromatic CKs – appear when an aromatic ring (either unsubstituted or substituted by a hydroxy or methoxy group) is attached to the N6 atom of adenine [26]. Substitution of the aromatic ring with a hydroxy group produces hydroxy derivatives such as ortho-topolin (oT) and meta-topolin (mT), while substitution with a methoxy group creates methoxy derivatives such as ortho-methoxytopolin (meoT) and meta-methoxytopolin (memT). Hydroxylation of the aromatic ring in meta position is a prerequisite for high CK activity [27, 28] and meta-topolin shows the most considerable effects in plants of all derivatives. The chemical structures of natural aromatic cytokinins are shown in Figure 2. Despite the isolation of the first naturally occurring aromatic CK as early as in the early 1970s [29], the distribution and function of aromatic CKs in the plant kingdom remain still unclear [30].
Figure 1.
Chemical structures of natural isoprenoid cytokinins.
Figure 2.
Chemical structures of natural aromatic cytokinins.
Synthetic CKs by chemical structure represent phenylurea derivatives. These substances show CK effects, however, to date they have not been discovered to be synthesized in plant tissues by normal metabolic pathways. The first identified synthetic CK was diphenylurea (DFU), discovered in 1955 [31]. The unexpected discovery of this compound stimulated the synthesis of numerous analogues such as CPPU (N-phenyl-N′- (2-chloro-4-pyridyl) urea) and TDZ (thidiazuron). These synthetic substances, presented in Figure 3, are highly stable and have stronger CK activity compared to zeatin [32, 33, 34, 35].
Figure 3.
Chemical structures of synthetic cytokinins.
1.2.1 Cytokinin catabolism
During plant development the levels of endogenous CKs are tightly metabolically regulated in plant tissues. The decreased amount of CKs is directly related to the increased activity of the catabolic enzyme, cytokinin oxidase/dehydrogenase (CKX, EC 1.5.99.12). This enzyme irreversibly degrades CKs by removing the N6-substituted side-chain to form adenine and the unsaturated aldehyde 3-methyl-2-butenal [36] (Figure 4). To date, CKX is the only known enzyme involved in the catabolism of specific CKs, and thus it represents a crucial factor in the control of CK levels in plant tissues. It is known that especially iP, tZ and their ribosides are sensitive to the CKX action. On the other hand, cZ is less sensitive [37] even though some CKX isoforms exhibit high affinity for the cZ isomer as well [38]. CK O-glucosides, DHZ, aromatic CKs and their derivatives are believed to be the non-substrates for CKX [39, 40, 41]. By contrast, Arabidopsis thaliana, barley and maize CKX isoforms showed ability to degrade CK N9-glucosides [37, 42, 43].
Figure 4.
Chemical reaction of irreversible cytokine degradation by CKX enzyme.
According to the chemical structure, CKX enzyme is a flavoprotein with the flavin adenosine dinucleotide (FAD) bound domain [44, 45]. The activity of CKX in plant tissues is induced by exogenously applied CKs, both CKX substrates and non-substrates [46, 47, 48]. The CKX activity was first discovered in tobacco tissue [49], and subsequently this enzyme has been shown widely distributed in numerous plant tissues such as e.g. maize [50], Vinca rosea tumor [36], Phaseolus vulgaris callus [51, 52], wheat [53], and tobacco and poplar calli [47, 54]. The CKX enzyme is encoded by a small family of genes whose number varies from species to species. In Arabidopsis thaliana, seven genes responsible for the synthesis of CKX enzyme (AtCKX1–7) have been identified. These genes are expressed variously in different plant tissues [55, 56]. Individual AtCKX genes are expressed in the same tissues where CKs are synthesized (axillary buds, trichomes and vascular tissues) or in the nearby of these tissues. Genes encoding the CKX enzyme have been cloned also from numerous plant species (a detailed literature review is presented in Table 1).
The list of CKX genes cloned and identified in different plant species.
1.3 Genetic transformation of plants using AtCKX genes
Genetic transformations of plants with specific genes encoding the CKX enzyme are beneficial for investigations of CK homeostasis. Thus, plants exhibiting increased overexpression of the CKX genes and a reduced amount of endogenous CKs can be obtained. Genetically engineered CKX plants can serve as a sound and valuable model system for studying physiological and morphological processes that are under control of CKs. To date, the impact of overexpression of different CKX genes on CK metabolism has been investigated in several plant species, as summarized in Table 2. The overexpression of CKX genes in tobacco and A. thaliana plants was demonstrated to affect significantly the phenotype of transformed plants, causing the cytokinin deficiency syndrome. In transgenic plants with this syndrome, apical dominance was decreased, shoots grew slowly, leaf size was reduced, the plants bloomed later and had a smaller number of flowers. Increased root growth size and activity of the root apical meristem (RAM) were observed. At the same time, decreased size and the shoot apical meristem (SAM) activity were also found. These symptoms were more evident in plants overexpressing AtCKX1 transgene than in those with AtCKX2 gene [55, 79]. As most of existing knowledge on regulation of CK levels and effects in plants have been obtained from studies of consequences of exogenously applied CKs, the specific CK-deficient transformants are expected to be more informative and to help in elucidation of the processes controlled by this group of phytohormones.
The list of plant species genetically transformed using CKX genes isolated from Arabidopsis thaliana.
In plant shoots, CKs stimulate function of apical meristem and cell divisions. The stimulatory role of CKs on cytokinesis was discovered a long time ago [85]. Recently, it has been confirmed by analyses of transgenic AtCKX A. thaliana plants with reduced endogenous CK levels. Ultrastructural analysis of SAM cells showed clear cytological changes indicating detention of cell division and accelerated cell differentiation in transgenic CK-deficient AtCKX A. thaliana plants [86]. The development of vascular elements is also regulated by CKs. The optimal concentration of CKs in combination with auxins affects the formation of phloem [87]. In the shoots of transgenic AtCKX A. thaliana plants, xylem and phloem were reduced [55]. In addition, CKs control leaf formation and growth. Overexpression of the CKX transgenes in AtCKX A. thaliana plants significantly reduced leaf surface, which is undoubtedly related to cell divisions as previously described [55]. It has been shown that CKs are not able to completely prevent, but can significantly delay the senescence process [88]. Thus, it can be assumed that a reduced amount of endogenous CKs in CK-deficient plants accelerates senescence. Despite these expectations, the leaves of transgenic AtCKX A. thaliana plants did not show accelerated senescence [55]. The role of CKs during the plant reproductive development was approved by transgenic AtCKX A. thaliana plants, which formed a smaller number of flowers [55]. Enlarged embryos have also been observed in these plants, indicating a role of CKs in cell division during embryogenesis. The enlargement of embryos could be explained on the basis of CKX genes overexpression, occurring only in the later stages of embryogenesis [89].
Unlike the shoots, CKs inhibit the function of the RAM. Using the transgenic A. thaliana AtCKX plants, it has been confirmed that reduced CK levels strengthen the root system. The primary root growth is directly related to intense cell divisions in the root meristem of transgenic A. thaliana AtCKX plants [55]. The results obtained in these transformants are consistent with those by [90] demonstrating that exogenously applied CKs inhibited root elongation of untransformed plants by reducing the RAM size as well as reducing cell divisions. One would expect that the plants transformed with CKX genes have inhibited root branching. However, CK-deficient transgenic AtCKX A. thaliana plants surprisingly developed numerous lateral roots that elongated rapidly [55].
1.4 Centaurium erythraea Rafn.
Common centaury, Centaurium erythraea Rafn. (syn. C. umbellatum Gillib and C. minus Moench), is the most known species from Gentianaceae family. Thanks to numerous pharmacological activities, centaury is listed as a medicinal plant species. In traditional medicine, centaury is used to treat febrile conditions, anaemia, jaundice, gout and to regulate blood sugar [91]. Bitter centaury juices have also been used for an increment of appetite, digestion stimulation and treatment of gastrointestinal tract diseases [92].
Nowadays, C. erythraea is rapidly disappearing from natural habitat and it is marked as an endangered plant species. Fortunately, even almost twenty years ago centaury showed vigorous regenerative potential in vitro [93, 94, 95, 96, 97]. The most interesting and essential information is that spontaneous morphogenesis of centaury is possible on nutrition medium without addition of any plant growth regulator [98]. In addition, during the years, centaury has also been used for studies of numerous developmental processes [99, 100, 101]. On the other hand, only few literature data exist describing genetic transformation of centaury. To date, most of Gentianaceae species, including C. erythraea, were usually genetically transformed only with A. rhizogenes [93, 102]. These investigations have mostly based and described the efficiency of plant regeneration and selection of transformed plant tissues. All of these previous investigations also encouraged us to use A. tumefaciens in genetic transformation of centaury for the first time [84]. The successful production of transgenic AtCKX centaury plants allowed to use them as a suitable model for studies of numerous physiological and developmental processes under endogenous phytohormonal control.
2. Methodology
All of the in vitro culture experiments were performed with plant material originated from C. erythraea Rafn. seeds, obtained from Jelitto Staudensamen GmbH, Schwarmstedt, Germany. The plant tissue culture methods were used to establish a solid centaury root culture further utilizable in the genetic transformation process. In all in vitro culture experiments, half-strength MS hormone-free medium (½MS) [103] solidified with 0.7% agar and supplemented with 3% sucrose and 100 mg l−1myo-inositol was used. The A. tumefaciens strain GV3101 harbouring the pBinHTX plasmid with either the AtCKX1 or AtCKX2 gene under the control of the 35S promoter with Triple X enhancer [79] was applied for the transformation of C. erythraea roots. Usual molecular biology analyses, including PCR and qPCR reactions, were performed to confirm the insertion of AtCKX transgenes into centaury genome. The CKX activity was determined by in vitro assays based on the conversion of [2-3H]iP (prepared by the Isotope Laboratory, Institute of Experimental Botany AS CR, Prague, Czech Republic) to [3H] adenine. Detection and quantification of endogenous phytohormones were performed using HPLC/MS system with TSQ Quantum Ultra AM triple-quadrupole high-resolution mass spectrometer (Thermo Electron Corp., San Jose, CA, U.S.A.) operated in the positive-SRM mode. Secondary metabolites including secoiridoids (swertiamarin and gentiopicrin) and xanthones (eustomin and demethyleustomin) were analysed on an Agilent series 1100 HPLC instrument with DAD detector and a reverse phase Zorbax SB-C18 analytical column. The secondary metabolite profile of transgenic centaury plants was determined to as certain whether these plants might serve as potential producers of anti-cancer compounds. The effect of non-transformed and transgenic AtCKX centaury methanol extracts on antibacterial and antifungal activity on bacteria and microfungi was also investigated. The salinity tolerance of non-transformed and AtCKX transgenic centaury shoots and roots to graded NaCl concentrations were tested separately in vitro. All plant material and methods used in investigations are presented in Table 3. The illustrative presentation of transgenic AtCKX centaury plants production in vitro and all further applied analyses are shown in Figure 5.
Plant material
Methods
References
C. erythraea Rafn. seeds
Establishment of a solid root culture by plant tissue culture methods
The list of all plant materials and methods used during the investigation of transgenic AtCKX centaury plants grown in vitro.
Figure 5.
Illustrative presentation of transgenic AtCKX centaury plants production in vitro and all subsequent investigations conducted to date.
3. Molecular and morphological characterisation of transgenic AtCKX centaury plants
The possibility of direct regeneration of centaury shoots and/or somatic embryos makes root cultures a good model for studying in vitro morphogenesis of this plant species. Root cultures, in addition to easy manipulation, are characterised by other advantages such as small differences in physiological responses as well as high growth potential and metabolic activity. During the time, it has been shown that root cultures are suitable for production of genetically, biochemically and phenotypically stable plant material without somaclonal variation. The plant cultures, including a callus phase, theoretically promote a higher mutation rate [108]. The first literature data describing centaury root culture appeared almost twenty years ago, when the adventitious buds regeneration was shown to be induced with callus formation on initial root explants [109]. Spontaneous regeneration of adventitious buds in solid root culture was described, for the first time, by Subotić et al. [94]. These results motivated us to use root tips growing on½MS hormone-free medium as initial explants for further spontaneous regeneration and propagation of centaury plants. In our investigations, a solid root culture has been originally proved to be also a suitable and grateful model for genetic transformation of this medicinal plant species using A. tumefaciens.
Because of prosperous selection efficiency and adequate regenerative capacity of the transformants, the selection of suitable explants for initial infection with A. tumefaciens represents the most crucial step for a successful process of genetic transformation [110]. The potentially transformed centaury shoots formed directly on the root explants, without callus induction and thus minimized possible somaclonal variation [84]. Genomic PCR with specific primers confirmed the successful integration of the AtCKX1 and AtCKX2 transgenes into centaury genome in 30% and 28.2% of the analysed hygromycin-resistant lines, respectively. The quantification of transgene expression was confirmed by highly sensitive and precise method, qRT-PCR. As transgenic AtCKX1 lines showed low expression and unspecific amplification of cDNA, optimization of PCR and qPCR protocols was necessary. The optimized protocol used for real-time quantification of the transgene expression included combining a gradient of annealing temperatures with the application of seven different PCR enhancers: formamide, DMSO, glycerol, ethylene glycol, trehalose, BSA and Tween-20 [111]. The expression of AtCKX1 and AtCKX2 transgenes was detected and quantified in 50% and 63.6% transgenic lines, respectively. The AtCKX1 roots showed slightly higher expression than shoots whereas the AtCKX2 transgene was better expressed in shoots compared to roots. Evidently, quantitative differences in transgene expression after incorporation into genome of different centaury lines depended on the transgene copy number and positional variations of transgenes.
All of the AtCKX transgenic centaury lines showed decreased regeneration capacity in root culture on the solid hormone-free ½MS medium in comparison to control. However, regenerated shoots spontaneously rooted on the same medium. On the other hand, fresh weight of shoots in most of analysed transgenic centaury lines showed no significant difference in comparison to control. In analogy, the root culture of Dendrobium orchid overexpressing DSCKX1 transgene proved reduced shoot regeneration with decreased biomass compared to the non-transformed controls [112]. Similarly, the increased CKX expression affected on shortened internodes and reduced leaf surface of transgenic Arabidopsis and tobacco shoots [44].
4. CKX activity and endogenous phytohormone content in transgenic AtCKX centaury plants
All centaury transgenic AtCKX1 and AtCKX2 shoots and roots showed increased CKX activity. Two to ten times higher CKX activity was detected in roots compared to shoots, which corresponds with previously demonstrated data in other plants such as tobacco [113], maize and barley [41, 64, 114]. In analogy to AtCKX-overexpressing Arabidopsis plants [55] the higher CKX activity was detected in shoots and roots of AtCKX2 than AtCKX1 centaury lines.
As the CKX represents a valuable regulatory enzyme in CK metabolism, CKX activity directly affects the CK amount in plant tissues [115, 116], the contents and profile of endogenous CKs in centaury non-transformed and AtCKX transgenic plants have been determined. In general, centaury shoots grown in vitro produced higher content of CKs than roots [104]. Considering production and dominant location of CKs in the roots this distribution of endogenous CKs may be a consequence of in vitro conditions itself. The roots of centaury plants overexpressing AtCKX transgenes showed higher content of total CKs in comparison to non-transformed ones [84]. Previous investigations on AtCKX overexpressing Arabidopsis and tobacco plants revealed that increased CKX activity affected on the reduction of different endogenous CK metabolites [44, 55, 79]. Theoretically, increased CKX activity should cause a decline in the total CKs amount. Regardless of this, overexpression of AtCKX genes did not contribute to lowered total CKs content in transgenic centaury plants but rather to an alteration in the spectrum of particular CK types. In analogy, transgenic potato plants overexpressing AtCKX genes were not found to show decreased total endogenous CK contents while bioactive CK forms were predominantly reduced [82, 83]. Accordingly, although enhanced CKX activity results in more substantial CK degradation in plant tissues it might simultaneously serve as an indirect signal for the plant cell to intensify the biosynthesis of CKs. It may lead to higher accumulation of total endogenous CKs and seems to be a mechanism of re-establishment and maintenance of CK homeostasis in plants [117].
Beside the total CKs contents, an altered amounts of individual endogenous CKs groups were specific forAtCKX transgenic centaury plants [104]. The reported considerable differences in endogenous CK spectra might reflect the distinct position and/or the number of transgene copies. The most specific characteristic of transgenic centaury plants was a remarkable reduction of bioactive CK forms, including free bases and their corresponding ribosides. Similarly, the AtCKX transgenic tobacco and potato plants also showed decreased contents of bioactive CKs [79, 82, 83, 118, 119]. In transgenic centaury plants, the reduction of bioactive CKs was organ-specific. Thus, considerably declined levels of bioactive CK forms were found in the AtCKX shoots although their CKX activity was not enhanced. On the other hand, a significant reduction of bioactive CKs levels was shown in AtCKX roots together with an increased CKX activity. A possible explanation is that CKs are probably more degraded in roots, the main location of biosynthesis, and accordingly, a smaller content of bioactive CKs is further transported to centaury shoots. Another one attribute of the AtCKX transgenic centaury plants was represented by increased amounts of CK storage forms (O-glucosides) and/or of irreversibly inactive (or weakly active)CK N-glucosides [104]. This is in accordance with recent literature data demonstrating the same CK pattern in AtCKX1 transgenic potato plants [83]. Simultaneously, the concentrations of CK precursors in transgenic centaury plants were higher than those of bioactive CKs, which also corresponds with the potato plants [82].
To summarize, the AtCKX transgenic centaury plants are characterized by altered CK profiles with reduced levels of bioactive CK forms and, at the same time, increased amounts of storage forms (CK O-glucosides), inactive (or weakly active) forms (CK N-glucosides) andCK nucleotides. Previous investigations showed that overexpression of AtCKX transgene increased the production of storage CK derivatives in transgenic potato plants [81, 82]. It can be assumed that introduction of AtCKX transgenes into centaury genome altered CK metabolism in a way leading to the increased production of endogenous CK conjugates. Simultaneously, the level of bioactive CK forms seems to be controlled, besides the CKX activity, by additional regulatory mechanisms involved in CK metabolism and transport throughout the plant tissues. Considering that bioactive CKs are the preferred substrates of CKX, it is also possible that plant tissues activate new CK biosynthetic pathways and further initiate conversion to storage compounds, deactivation forms and/or CK nucleotides.
Numerous physiological and developmental processes in plants are regulated by the cooperation of CKs and auxins [120, 121, 122, 123]. Overexpression of AtCKX transgenes in centaury plants altered not only the CK homeostasis, but at the same time it affected metabolism of auxins. In transgenic centaury plants, the auxin indole-3-acetic acid (IAA) concentration was significantly decreased compared to the control. In analogy, previous literature data uncovered that an enhancement of the CKX activity led to the decline of endogenous CK levels and simultaneously to the reduction of endogenous IAA content in A. thaliana [79, 122]. In contrast, increased IAA concentration was reported in the AtCKX1-overexpressing potato plants [83]. These findings support the fact that the balance between CKs and auxins represents the most crucial factor in the shoot and root development in plants [55, 79].
In transgenic AtCKX centaury roots, the IAA/bioactive CKs ratio was lowered in comparison to control. It might be a consequence of a reduced regeneration capacity of transgenic centaury roots [84]. Considering that endogenous IAA content was lowered in the roots, it is presumable that AtCKX-overexpression weakened the regeneration of shoots in solid roots culture. In transgenic centaury shoots, the IAA/bioactive CKs ratios varied considerably. Contrary to the centaury plants, endogenous IAA content was significantly increased in the shoots and roots of transgenic AtCKX1 potato [83].
An altered CK metabolism resulting from the overexpression of AtCKX transgenes definitely affected IAA content and IAA/bioactive CKs ratio in transformed centaury plants. Different biosynthetic capacity for CK and IAA production could be a reason for the organ-specific differences among AtCKX transgenic plants.
5. Secondary metabolites in transgenic AtCKX centaury plants
It is known that in vitro cultivation, although under controlled temperature and light conditions, stimulates increased secondary metabolites accumulation [124]. The previous investigations also showed that genetic transformation of centaury roots using A. rhizogenes affected the production of secondary metabolites [102, 125]. Considering that centaury plants were transformed using A. tumefaciens carrying AtCKX genes for the first time, it was interesting to reveal the effects of altered levels of endogenous CKs on secondary metabolites production of these transgenic centaury plants. The results of our analyses indicated modifications of the secondary metabolites production in this valuable medicinal plant species as a consequence of the AtCKX transgenes overexpression [105]. In methanol extracts of AtCKX transgenic centaury plants, the presence of bitter secoiridoids (swertiamarin and gentiopicrin) and xanthones (eustomin and demethyleustomin) was detected. The fact that AtCKX transgenic centaury plants produced the same compounds as non-transformed ones confirmed that there were no qualitative differences in specific secondary metabolites resulting from the AtCKX overexpression. On the other hand, quantitative changes in the secondary metabolite contents were found. Considering that gentiopicrin originates from swertiamarin in the metabolic pathway of iridoids, it is possible that increased activity of the swertiamarin → gentiopicrin converting enzyme stimulated gentiopicrin production. In all AtCKX transgenic centaury shoots, secoiridoid swertiamarin represented the predominant component. Still, in almost all centaury AtCKX shoots the content of swertiamarin was lowered in comparison to control shoots from in vitro cultures as well as from natural habitat. On the other hand, the production of the second bitter secoiridoid, gentiopicrin, was increased in the majority of AtCKX transgenic centaury shoots compared to control ones, grown in vitro and collected in natural habitat.
Unlike shoots, the AtCKX transgenic centaury roots produced decreased content of both secoiridoids, swertiamarin and gentiopicrin, in comparison to control roots grown in vitro and from natural habitat. It was shown previously that swertiamarin dominated in centaury shoots and roots from natural habitat while gentiopicrin prevailed in centaury shoots and roots grown in vitro [126, 127, 128, 129].
Beside secoiridoids, the differences in xanthone content were also detected in AtCKX transgenic centaury plants. Most of AtCKX transgenic shoots and roots produced more eustomin and demethyleustomin than shoots from natural habitat but less than control in vitro shoots. The shoots and roots of only one transgenic centaury line, AtCKX1–29, produced significantly increased amount of both xanthones in comparison to both controls, shoots grown in vitro and those collected from natural habitat. It is important to note that shoots and roots of this transgenic line were characterised by a reduced level of bioactive forms of CKs [104].
5.1 Transgenic AtCKX centaury plants as potential producers of anti-cancer compounds and antimicrobials
The methanol extracts of in vitro cultured AtCKX1–29 transgenic centaury plants were also tested for their antimicrobial effects [106]. Four Gram-positive, four Gram-negative bacteria and eight species of microfungi were selected for these in vitro investigations. The methanol extracts of non-transformed centaury plants, as well as pure secoiridoids (swertiamarin and gentiopicrin) and xanthones (eustomin and demethyleustomin), were used as a control. In general, the extracts of all tested centaury shoots and roots showed an adequate antibacterial activity on all tested bacteria. Methanol extracts of non-transformed and AtCKX1–29 transgenic centaury roots and pure gentiopicrin had a low antibacterial activity on Micrococcus flavus, Escherichia coli and Enterobacter cloacae. Staphylococcus aureus was found the most sensitive bacterial species in the in vitro assays. It was also shown that the extract of non-transformed centaury shoots was more effective on S. aureus than antibiotics, streptomycin and ampicillin. Interestingly, the pure compounds such as swertiamarin, gentiopicrin, eustomin and demethyleustomin exhibited higher antibacterial activity on all tested bacteria than centaury methanol extracts and commercial antibiotics (streptomycin and ampicillin) used as a positive control. High antimicrobial activity could be ascribed to bitter secoiridoid glycosides, which corresponds with previous literature data [130, 131]. Beside antibacterial impact, all tested methanol extracts of centaury shoots and roots also showed antifungal effects. Most of the applied compounds proved high antifungal activities. All pure secoiridoids and xanthones were more effective than methanol extracts against all fungi. In general, xanthone eustomin showed even 100 times higher antifungal potential than both of the applied mycotics (ketoconazole and bifonazole) used as a positive control. These results are significant as they represent the first report describing centaury xanthones as potential antimicrobials. In general, methanol extracts of non-transformed and transgenic AtCKX1 centaury shoots and roots showed better antibacterial activity, while pure secoiridoids and xanthones were more active against fungi.
Antioxidant properties of centaury plants and pure xanthones were reported almost twenty years ago [132, 133]. Considering that transgenic AtCKX1–29 roots produced increased amounts of xanthones, their potential antioxidant activity was evaluated [106]. The transgenic AtCKX1 centaury shoots and roots were found to exert higher antioxidant activity compared to non-transformed plants. The root extracts were two to five times more effective than those from the shoots. Transgenic AtCKX1–29 roots, containing the highest amounts of total phenolics, were the most effective the scavenging of the DPPH radicals. The antioxidant effects of transgenic centaury methanol extracts could be assigned to elevated levels of xanthone compounds. Accordingly, centaury plants with increased content of secondary metabolites could be of practical importance in developing novel drugs with a potential use in agronomy, veterinary, medicine and food industry.
6. Transgenic AtCKX centaury plants under salinity stress in vitro
Taking into account that centaury plants can inhabit saline soils in natural habitats and because it is known that CKs play an essential role in the salinity stress response, the effect of NaCl-induced stress on regeneration potential of centaury shoots and roots was evaluated. The investigated non-transformed as well as AtCKX1 and AtCKX2 transgenic lines showed different salinity tolerance to graded NaCl concentrations. In general, a higher salinity tolerance was found for roots compared to shoots. Furthermore, elevated NaCl concentrations in the culture medium had no inhibitory effect on centaury shoot growth. All centaury shoots, regenerated on media supplemented with different NaCl concentrations, showed similar morphology as shoots regenerated in solid root cultures grown on NaCl-free media. Similarly, Šiler et al. [134] demonstrated that centaury plants keep the rosette forms under salt stress in vitro. Variations in the salt tolerance could be explained by differences among centaury genotypes, which is related to the recent report describing this species as a common salt-tolerant medicinal herb from Mediterranean region [135]. Whereas the non-transformed and AtCKX1 transgenic line showed the same trend of shoot regeneration potential under salt stress conditions, the AtCKX2 transgenic lines differed exhibiting a gradually decreased frequency of regeneration, the average number of regenerated shoots and fresh shoot weight with increased NaCl in the medium. The AtCKX transgenic centaury lines differed in their salinity tolerance from the non-transformed control indicating possible involvement of CKs in this process.
Beside morphological traits, biochemical characterisation of AtCKX transgenic centaury plants was determined. Amount of endogenous proline is considered one of the factors involved in the plant stress tolerance. Accordingly, an increase in the proline content was found in all AtCKX centaury shoots and roots grown on graded NaCl concentrations. Evaluation of malondialdehyde (MDA) level and hydrogen peroxide (H2O2) in centaury plants grown in vitro during salt stress also provided evidence of their oxidative stress tolerance. In all centaury AtCKX shoots and roots, MDA and H2O2 contents were increased at graded NaCl concentrations. These findings correspond well with literature data demonstrating an enhancement of MDA and H2O2 contents under salt stress [136, 137].
On the other hand, AtCKX transgenic centaury plants showed altered reactive oxygen species (ROS) homeostasis. It is rather difficult to summarize and conclude how salt stress affected on antioxidative enzymes activity. An increase of the superoxide dismutase (SOD) activity was detected in all centaury plants grown on graded NaCl concentrations. Interestingly, the SOD activity was always higher in roots than in shoots. Enhanced catalase (CAT) and peroxidase (POX) activities were detected in centaury plants grown on graded NaCl concentrations as well. Similarly to centaury, transgenic AtCKX2 tobacco plants showed increased SOD and CAT activity in comparison to wild type while lower SOD activity was found in roots than in shoots under salt stress ex vitro [118]. Increased SOD, CAT and POX activities in salt-stressed non-transformed and AtCKX transgenic centaury plants indicates a crucial role of these antioxidant enzymes in protecting plant tissues from superoxide radical and hydrogen peroxide and suggests their association with improved tolerance of plants to environmental stress conditions.
Evidently, the salinity stress caused morphological, physiological and biochemical changes in both non-transformed and AtCKX transgenic centaury plants. Elevated proline, MDA and H2O2 contents as well as increased antioxidative enzymes activities confirmed that centaury plants are trying to overcome salt stress conditions to maintain or re-establish their normal growth and development.
7. Conclusion and future perspectives
This chapter presents a survey of obtaining and developing transgenic AtCKX centaury plants grown in vitro and is intended to provide a synthesis of the existing literature data. In addition, recent achievements in the characterisation of transgenic centaury plants and determination as well as application of their secondary metabolite content is reported in details.
Centaury is a medicinal plant species endangered in natural habitat. A successful protocol for its genetic transformation using A. tumefaciens to introduce foreign, AtCKX, genes is described here. This protocol favours centaury root tips as primary infection explants that can directly regenerate shoots without callus phase. As the in vitro regeneration of centaury is possible without the addition of exogenous plant growth regulator(s), in vitro culture represents an optimal model for studying metabolic pathways of endogenous phytohormones including CKs without any interferences from other exogenous hormones present in nutrition medium. The first step of a successful genetic transformation of centaury was the establishment of stable transformants with AtCKX overexpression and enhanced CKX activity. Molecular, morphometric and developmental characterisation of obtained AtCKX-transgenic plants as well as chemical profiling of CK metabolites represented the following steps. The subsequent secondary metabolite analyses showed antibacterial and antifungal activity of transgenic centaury plants that can also be used as potential producers of anti-cancer compounds. Considering that centaury can inhabit saline soils in natural habitats, salinity tolerance of transgenic AtCKX centaury plants was investigated as well.
All obtained results were summarized and indicated that transgenic AtCKX centaury plants can serve as a suitable material for investigations of numerous physiological and developmental processes that are under endogenous phytohormonal control. Also, further studies of endogenous plant hormone regulations during NaCl-induced stress might represent an useful tool for better understanding the salinity tolerance of non-transformed and CK-deficient transgenic centaury plants grown in vitro. Future research will certainly be focused on the identification of centaury candidate genes specifically expressed during the salt stress to understand complex regulatory pathways in response of centaury plants to salinity in vitro.
Acknowledgments
This work was supported by the Ministry of Education, Science and Technological Development of the Republic of Serbia, contract number 451-03-68/2020-2114/200007, and by the Czech Science Foundation, project number 19-12262S.
Conflict of interest
The authors declare that they have no conflict of interest.
Abbreviations
CAT
catalase
CKs
cytokinins
CKX
cytokinin oxidase/dehydrogenase
DPPH
2,2-diphenyl-1-picryl-hydrazyl-hydrate
HPLC
high-performance liquid chromatography
MDA
malondialdehyde
MS
Murashige and Skoog medium
MS spectrometry
mass spectrometry
POX
peroxidase
ROS
reactive oxygen species
SOD
superoxide dismutase
\n',keywords:"centaury, Centaurium erythraea Rafn., in vitro propagation, AtCKX genes, cytokinins, salinity",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/74053.pdf",chapterXML:"https://mts.intechopen.com/source/xml/74053.xml",downloadPdfUrl:"/chapter/pdf-download/74053",previewPdfUrl:"/chapter/pdf-preview/74053",totalDownloads:135,totalViews:0,totalCrossrefCites:0,dateSubmitted:"September 8th 2020",dateReviewed:"October 28th 2020",datePrePublished:"December 14th 2020",datePublished:null,dateFinished:"November 13th 2020",readingETA:"0",abstract:"The production and breeding of plants with desired properties are possible by a fundamental biotechnological technique, genetic engineering. Applying and developing of genetic engineering procedures also enable preservation and improvement of plant species endangered in nature, including medicinal plant common centaury (Centaurium erythraea Rafn.). Numerous developmental processes in plants are controlled by cytokinins (CKs). The only so far known enzyme involved in CK catabolism is cytokinin oxidase/dehydrogenase (CKX). Genes coding for two Arabidopsis CKX isoforms, AtCKX1 and AtCKX2, were successfully introduced into centaury root explants. Subsequently, the contents of endogenous CKs in AtCKX-overexpressing centaury plants grown in vitro were investigated. Simultaneous secondary metabolite analyses showed antibacterial and antifungal activity of transgenic centaury plants and suggested their use as potential producers of anti-cancer compounds. Considering that centaury can inhabit saline soils in natural habitats, following investigations included evaluation of salinity tolerance in vitro. All obtained and summarized results indicated that transgenic AtCKX centaury plants could serve as a suitable model for studies of numerous physiological and developmental processes under endogenous phytohormonal control.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/74053",risUrl:"/chapter/ris/74053",signatures:"Milana Trifunović-Momčilov and Václav Motyka",book:{id:"10365",title:"Plant Breeding - Current and Future Views",subtitle:null,fullTitle:"Plant Breeding - Current and Future Views",slug:null,publishedDate:null,bookSignature:"Dr. Ibrokhim Y. Abdurakhmonov",coverURL:"https://cdn.intechopen.com/books/images_new/10365.jpg",licenceType:"CC BY 3.0",editedByType:null,isbn:"978-1-83968-310-7",printIsbn:"978-1-83968-309-1",pdfIsbn:"978-1-83968-311-4",editors:[{id:"213344",title:"Dr.",name:"Ibrokhim Y.",middleName:null,surname:"Abdurakhmonov",slug:"ibrokhim-y.-abdurakhmonov",fullName:"Ibrokhim Y. Abdurakhmonov"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:null,sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_1_2",title:"1.1 Genetic transformations of plants",level:"2"},{id:"sec_1_3",title:"1.1.1 Plasmids of bacteria of the genus Agrobacterium",level:"3"},{id:"sec_2_3",title:"1.1.2 Mechanism of genetic transformation of a plant cell using Agrobacterium tumefaciens",level:"3"},{id:"sec_4_2",title:"1.2 Cytokinins",level:"2"},{id:"sec_4_3",title:"Table 1.",level:"3"},{id:"sec_6_2",title:"1.3 Genetic transformation of plants using AtCKX genes",level:"2"},{id:"sec_7_2",title:"1.4 Centaurium erythraea Rafn.",level:"2"},{id:"sec_9",title:"2. Methodology",level:"1"},{id:"sec_10",title:"3. Molecular and morphological characterisation of transgenic AtCKX centaury plants",level:"1"},{id:"sec_11",title:"4. CKX activity and endogenous phytohormone content in transgenic AtCKX centaury plants",level:"1"},{id:"sec_12",title:"5. Secondary metabolites in transgenic AtCKX centaury plants",level:"1"},{id:"sec_12_2",title:"5.1 Transgenic AtCKX centaury plants as potential producers of anti-cancer compounds and antimicrobials",level:"2"},{id:"sec_14",title:"6. Transgenic AtCKX centaury plants under salinity stress in vitro",level:"1"},{id:"sec_15",title:"7. Conclusion and future perspectives",level:"1"},{id:"sec_16",title:"Acknowledgments",level:"1"},{id:"sec_19",title:"Conflict of interest",level:"1"},{id:"sec_18",title:"Abbreviations",level:"1"}],chapterReferences:[{id:"B1",body:'Slater A, Scott NW, Fowler MR. Plant Biotechnology. The Genetic Manipulation of Plants. New York: Oxford University Press; 2004. pp. 35-52'},{id:"B2",body:'Shou H, Frame BR, Whitham SA, Wang K. Assessment of transgenic maize events produced by particle bombardment or Agrobacterium-mediated transformation. 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Quantification of arabinogalactan proteins during in vitro morphogenesis induced by β-D-glucosyl Yariv reagent in Centaurium erythraearoot culture. Acta Physiologiae Plantarum. 2014;36:1187-1195. DOI: 10.1007/s11738-014-1495-y'},{id:"B100",body:'Filipović BK, Simonović AD, Trifunović MM, Dmitrović SS, Savić JM, Jevremović SB, et al. Plant regeneration in leaf culture of Centaurium erythraea Rafn. Part 1: The role of antioxidant enzymes. Plant Cell Tissue and Organ Culture. 2015;121:703-719. DOI: 10.1007/s11240-015-0740-4'},{id:"B101",body:'Simonović AD, Filipović BK, Trifunović MM, Malkov SN, Milinković VP, Jevremović SB, et al. Plant regeneration in leaf culture of Centaurium erythraea Rafn. Part 2: The role of arabinogalactan proteins. Plant Cell Tissue and Organ Culture. 2015;21:721-739. DOI: 10.1007/s11240-020-01801-w'},{id:"B102",body:'Piatczak E, Krolicka A, Wysokinska H. Genetic transformation of Centaurium erythraea Rafn by Agrobacterium rhizogenes and the production of secoiridoids. Plant Cell Reports. 2006;25:1308-1315'},{id:"B103",body:'Murashige T, Skoog F. A revised medium for rapid growth and bioassays with tobacco tissue cultures. Physiologia Plantarum. 1962;15:473-479'},{id:"B104",body:'Trifunović M, Motyka V, Cingel A, Subotić A, Jevremović S, Petrić M, et al. Changes in cytokinin content and altered cytokinin homeostasis in AtCKX1 and AtCKX2-overexpressing centaury (Centaurium erythraea Rafn.) plants grown in vitro. Plant Cell Tissue and Organ Culture. 2015;120:767-777'},{id:"B105",body:'Trifunović-Momčilov M, Krstić-Milošević D, Trifunović S, Podolski-Renić A, Pešić M, Subotić A (2016) Secondary metabolite profile of transgenic centaury (Centaurium erythraea Rafn.) plants, potential producers of anticancer compounds. Transgenesis and Secondary Metabolism, Reference Series in Phytochemistry. S Jha (ed), Springer International Publishing, DOI 10.1007/978-3-319-27490-4_5-1. ISBN: 978-3-319-27490-4'},{id:"B106",body:'Trifunović-Momčilov M, Krstić-Milošević D, Trifunović S, Ćirić A, Glamočlija J, Jevremović S, et al. Antimicrobial activity, antioxidant potential and total phenolic content of transgenic AtCKX1 centaury (Centaurium erythraea Rafn.) plants grown in vitro. Environmental Engineering and Management Journal. 2019;18:2063-2072'},{id:"B107",body:'Trifunović-Momčilov M, Paunović D, Milošević S, Marković M, Jevremović S, Dragićević IČ, et al. Salinity stress response of non-transformed and AtCKX transgenic centaury (Centaurium erythraea Rafn.) shoots and roots grown in vitro. Annals of Applied Biology. 2020;177:74-89'},{id:"B108",body:'Krishna H, Alizadeh M, Singh D, Singh U, Chauhan N, Eftekhari M, et al. Somaclonal variations and their applications in horticulturalcrops improvement. 3 Biotech. 2016;6:54'},{id:"B109",body:'Piatczak E, Wysokinska H. In vitro regeneration of Centaurium erythraea Rafn from shoot tips and other seedling explants. Acta Societatis Botanicorum Poloniae. 2003;72:283-288'},{id:"B110",body:'Kim JB. Development of efficient regeneration and transformation systems in Alstroemeria. PhD thesis. The Netherlands: Wageningen University; 2005. pp. 1-100'},{id:"B111",body:'Simonović A, Trifunović M, Raspor M, Cingel A, Bogdanović M, Dragićević M, et al. Dimethyl sulfoxide improves sensitivity and specificity of RT-PCR and qRT-PCR amplification of low-expressed transgenes. Archives of Biological Sciences. 2012;64:865-876'},{id:"B112",body:'Yang SH, Yu H, Xu Y, Goh YC. Investigation of cytokinin-deficient phenotypes in Arabidopsis by ectopic expression of orchid DSCKX1. FEBS Letters. 2003;555:291-296'},{id:"B113",body:'Motyka V, Faiss M, Strnad M, Kamínek M, Schmülling T. Changes in cytokinin content and cytokinin oxidase activity in response to derepression of ipt gene transcription in transgenic tobacco calli and plants. Plant Physiology. 1996;112:1035-1043'},{id:"B114",body:'Jones RJ, Schreiber BMN. Role and function of cytokinin oxidase in plants. Journal of Plant Growth Regulation. 1997;23:123-134'},{id:"B115",body:'Frébort I, Kowalska M, Hluska T, Frébortová J, Galuszka P. Evolution of cytokinin biosynthesis and degradation. Journal of Experimental Botany. 2011;62:2431-2452'},{id:"B116",body:'Kieber JJ, Schaller GE. Cytokinins. Arabidopsis book. 2014;12:e0168'},{id:"B117",body:'Kamínek M, Motyka V, Vaňková R. Regulation of cytokinin content in plant cells. Physiologia Plantarum. 1997;101:689-700'},{id:"B118",body:'Mytinova Z, Motyka V, Haisel D, Gaudinová A, Lubovská Z, Wilhemová N. Effect of abiotic stresses on the activity of antioxidative enzymes and contents of phytohormones in wild type and AtCKX2 transgenic tobacco plants. Biologia Plantarum. 2010;54:461-470'},{id:"B119",body:'Macková H, Hronková M, Dobrá J, Turečková V, Novák O, Lubovská Z, et al. Enhanced drought and heat stress tolerance of tobacco plants with ectopically enhanced cytokinin oxidase/dehydrogenase gene expression. Journal of Experimental Botany. 2013;64:2805-2815'},{id:"B120",body:'Eklöff S, Astot C, Blackwell J, Mortiz T, Olsson O, Sandberg G. Auxin-cytokinin interactions in wild-type and transgenic tobacco. Plant & Cell Physiology. 1997;38:225-235'},{id:"B121",body:'Nordström A, Tarkowski P, Tarkowska D, Norbaek R, Astot C, Doležal K, et al. Auxin regulation of cytokinin biosynthesis in Arabidopsis thaliana: a factor of potential importance for auxin-cytokinin-regulated development. Proc Natl Acad Sci USA. 2004;101:8039-8044'},{id:"B122",body:'Jones B, Andersson Gunnears S, Petrsson SV, Tarkowski P, Graham N, May S, et al. Cytokinin regulation of auxin synthesis in Arabidopsis involves a homeostatic feedback loop regulated via auxin and cytokinin signal transduction. Plant Cell. 2010;22:2956-2969'},{id:"B123",body:'Liu J, Mehdi S, Topping J, Tarkowski P, Lindsey K. Modelling and experimental analysis of hormonal crosstalk in Arabidopsis. Mol Systems Biol 6. Article number. 2010;373:1-13'},{id:"B124",body:'Hussain MS, Fareed S, Ansari S, Rahman MA, Ahmad IZ, Saeed M. Current approaches toward production of secondary plant metabolites. Journal of Pharmacy & Bioallied Sciences. 2012;4:10-20'},{id:"B125",body:'Janković T, Krstić D, Šavikin-Fodulović K, Menković N, Grubišić D. Xanthones and secoiridoids from hairy root cultures of Centaurium erythraea and C. pulchellum. Planta Med. 2002;68:944-946'},{id:"B126",body:'Van der Sluis WG (1985) Secoiridoids and xanthones in the genus Centaurium Hill (Gentianaceae) – a pharmacognostical study. Drukkerij Elinkwijk bv – Utrecht'},{id:"B127",body:'Van der Sluis WG. Chemotaxonomical investigations of the Genera Blackstonia and Centaurium (Gentianaceae). Plant Systematics and Evolution. 1985;149:253-286'},{id:"B128",body:'Janković T, Krstić D, Šavikin-Fodulović K, Menković N, Grubišić D. Comparative investigation of secoiridoid compounds of Centaurium erythraea grown in nature and cultured in vitro. Pharm and Pharmacol Lett. 1997;7:30-32'},{id:"B129",body:'Piatczak E, Wielanek M, Wysokinska H. Liquid culture system for shoot multiplication and secoiridoid production in micropropagated plants of Centaurium erythraea Rafn. Plant Science. 2005;168:431-437'},{id:"B130",body:'Šiler B, Mišić D, Nestorović J, Banjanac T, Glamočlija J, Soković M, et al. Antibacterial and antifungal screening of Centaurium pulchellum crude extracts and main secoiridoid compounds. Natural Products Communications. 2010;5:1525-1530'},{id:"B131",body:'Šiler B, Živković S, Banjanac T, Cvetković J, Nestorović-Živković J, Ćirić A, et al. Centauries as underestimated food additives: Antioxidant and antimicrobial potential. Food Chemistry. 2014;147:367-376'},{id:"B132",body:'Valentão P, Fernandes E, Carvalho F, Andrade PB, Seabra RM, Bastos ML. Antioxidant activity of Centaurium erythraeainfusion evidenced by its superoxide radical scavenging and xanthine oxidase inhibitory activity. Journal of Agricultural and Food Chemistry. 2001;49:3476-3479'},{id:"B133",body:'Pinto MMM, Sousa ME, Nascimento MSJ. Xanthone derivatives: new insights in biological activities. Current Medicinal Chemistry. 2005;12:2517-2538'},{id:"B134",body:'Šiler B, MišićD FB, Popović Z, Cvetić T, Mijović A. Effects of salinity on in vitro growth and photosynthesis of common centaury (Centaurium erythraea Rafn.). Archives of Biological Sciences. 2007;59:129-134'},{id:"B135",body:'Petropoulos SA, Karkanis A, Martins N, Ferreira ICFR. Halophytic herbs of the Mediterranean basin: an alternative approach to health. Food and Chemical Toxicology. 2018;114:155-169'},{id:"B136",body:'Palma F, López-Gómez M, Tejera NA, Lluch C. Involvement of abscisic acid in the response of Medicago sativa plants in symbiosis with Sinorhizobium meliloti to salinity. Plant Science. 2014;223:16-24'},{id:"B137",body:'Ikbal FE, Hernández JA, Barba-Espín G, Koussa T, Aziz A, Faize M, et al. Enhanced salt-induced antioxidative responses involve a contribution of polyamine biosynthesis in grapevine plants. Journal of Plant Physiology. 2014;171:779-788'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Milana Trifunović-Momčilov",address:"milanag@ibiss.bg.ac.rs",affiliation:'
Department for Plant Physiology, Institute for Biological Research “Siniša Stanković” – National Institute of Republic of Serbia, University of Belgrade, Bulevar despota Stefana 142, Serbia
Institute of Experimental Botany of the Czech Academy of Sciences, Rozvojová 263, Czech Republic
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IntechOpen’s team of Scientific Advisors supports the publishing team by providing editorial and academic input and ensuring the highest quality output of free peer-reviewed articles. The Boards consist of independent external collaborators who assist us on a voluntary basis. Their input includes advising on new topics within their field, proposing potential expert collaborators and reviewing book publishing proposals if required. Board members are experts who cover major STEM and HSS fields. All are trusted IntechOpen collaborators and Academic Editors, ensuring that the needs of the scientific community are met.
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Physical Sciences, Technology and Engineering Board
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Chemistry
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Ayben Kilislioglu - Department of Chemical Engineering Istanbul University, İstanbul, Turkey
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Goran Nikolic - Faculty of Technology, University of Nis, Leskovac, Serbia
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Mark T. Stauffer - Associate Professor of Chemistry, The University of Pittsburgh, USA
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Margarita Stoytcheva - Autonomous University of Baja California Engineering Institute Mexicali, Baja California, Mexico
Joao Luis Garcia Rosa - Associate Professor Bio-inspired Computing Laboratory (BioCom) Department of Computer Science University of Sao Paulo (USP) at Sao Carlos, Brazil
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Jan Valdman - Institute of Mathematics and Biomathematics, University of South Bohemia, České Budějovice, Czech Republic Institute of Information Theory and Automation of the ASCR, Prague, Czech Republic
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Earth and Planetary Science
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Jill S. M. Coleman - Department of Geography, Ball State University, Muncie, IN, USA
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İbrahim Küçük Erciyes - Üniversitesi Department of Astronomy and Space Sciences Melikgazi, Kayseri, Turkey
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Pasquale Imperatore - Electromagnetic Environmental Sensing (IREA), Italian National Council of Research (CNR), Naples, Italy
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Mohammad Mokhtari - Director of National Center for Earthquake Prediction International Institute of Earthquake Engineering and Seismology (IIEES), Tehran, Iran
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Engineering
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Narottam Das - University of Southern Queensland, Australia
\\n\\t
Jose Ignacio Huertas - Energy and Climate Change Research Group; Instituto Tecnológico y Estudios Superiores de Monterrey, Mexico
Likun Pan - Engineering Research Center for Nanophotonics and Advanced Instrument, Ministry of Education, Department of Physics, East China Normal University, China
\\n\\t
Mukul Chandra Paul - Central Glass & Ceramic Research Institute Jadavpur, Kolkata, India
\\n\\t
Stephen E. Saddow - Electrical Engineering Department, University of South Florida, USA
\\n\\t
Ali Demir Sezer - Marmara University, Faculty of Pharmacy, Department of Pharmaceutical Biotechnology, İstanbul, Turkey
\\n\\t
Krzysztof Zboinski - Warsaw University of Technology, Faculty of Transport, Warsaw, Poland
\\n
\\n\\n
Materials Science
\\n\\n
\\n\\t
Vadim Glebovsky - Senior Researcher, Institute of Solid State Physics, Chernogolovka, Russia Expert of the Russian Fund for Basic Research, Moscow, Russia
\\n\\t
Jianjun Liu - State Key Laboratory of High Performance Ceramics and Superfine Microstructure of Shanghai Institute of Ceramics, Chinese Academy of Sciences, China
\\n\\t
Pietro Mandracci - Department of Applied Science and Technology, Politecnico di Torino, Italy
\\n\\t
Waldemar Alfredo Monteiro - Instituto de Pesquisas Energéticas e Nucleares Materials Science and Technology Center (MSTC) São Paulo, SP, Brazil
Toshio Ogawa - Department of Electrical and Electronic Engineering, Shizuoka Institute of Science and Technology, Toyosawa, Fukuroi, Shizuoka, Japan
\\n
\\n\\n
Mathematics
\\n\\n
\\n\\t
Paul Bracken - Department of Mathematics University of Texas, Edinburg, TX, USA
\\n
\\n\\n
Nanotechnology and Nanomaterials
\\n\\n
\\n\\t
Muhammad Akhyar - Farrukh Nano-Chemistry Lab. Registrar, GC University Lahore, Pakistan
\\n\\t
Khan Maaz - Chinese Academy of Sciences, China & The Pakistan Institute of Nuclear Science and Technology, Pakistan
\\n
\\n\\n
Physics
\\n\\n
\\n\\t
Izabela Naydenova - Lecturer, School of Physics Principal Investigator, IEO Centre College of Sciences and Health Dublin Institute of Technology Dublin, Ireland
\\n\\t
Mitsuru Nenoi - National Institute of Radiological Sciences, Japan
\\n\\t
Christos Volos - Physics Department, Aristotle University of Thessaloniki, Greece
\\n
\\n\\n
Robotics
\\n\\n
\\n\\t
Alejandra Barrera - Instituto Tecnológico Autónomo de México, México
\\n\\t
Dusan M. Stipanovic - Department of Industrial and Enterprise Systems Engineering, University of Illinois at Urbana-Champaign
\\n\\t
Andrzej Zak - Polish Naval Academy Faculty of Navigation and Naval Weapons Institute of Naval Weapons and Computer Science, Gdynia, Poland
Petr Konvalina - Faculty of Agriculture, University of South Bohemia in České Budějovice, Czech Republic
\\n
\\n\\n
Biochemistry, Genetics and Molecular Biology
\\n\\n
\\n\\t
Chunfa Huang - Saint Louis University, Saint Louis, USA
\\n\\t
Michael Kormann - University Children's Clinic Department of Pediatrics I, Pediatric Infectiology & Immunology, Translational Genomics and Gene Therapy in Pediatrics, University of Tübingen, Tübingen, Germany
\\n\\t
Bin WU - Ph.D. HCLD Scientific Laboratory Director, Assisted Reproductive Technology Arizona Center for Reproductive Endocrinology and Infertility Tucson, Arizona , USA
\\n
\\n\\n
Environmental Sciences
\\n\\n
\\n\\t
Juan A. Blanco - Senior Researcher & Marie Curie Research Fellow Dep. Ciencias del Medio Natural, Universidad Publica de Navarra Campus de Arrosadia, Pamplona, Navarra, Spain
\\n\\t
Mikkola Heimo - University of Eastern Finland, Kuopio, Finland
\\n\\t
Bernardo Llamas Moya - Politechnical University of Madrid, Spain
\\n\\t
Toonika Rinken - Department of Environmental Chemistry, University of Tartu, Estonia
\\n
\\n\\n
Immunology and Microbiology
\\n\\n
\\n\\t
Dharumadurai Dhanasekaran - Department of Microbiology, School of Life Sciences, Bharathidasan University, India
Isabel Gigli - Facultad de Agronomia-UNLPam, Argentina
\\n\\t
Milad Manafi - Department of Animal Science, Faculty of Agricultural Sciences, Malayer University, Malayer, Iran
\\n\\t
Rita Payan-Carreira - Universidade de Trás-os-Montes e Alto Douro, Departamento de Zootecnia, Portugal
\\n
\\n\\n
Medicine
\\n\\n
\\n\\t
Mazen Almasri - King Abdulaziz University, Faculty of Dentistry Jeddah, Saudi Arabia Dentistry
\\n\\t
Craig Atwood - University of Wisconsin-Madison, USA Stem Cell Research, Tissue Engineering and Regenerative Medicine
\\n\\t
Oreste Capelli - Clinical Governance, Local Health Authority, Modena, Italy Public Health
\\n\\t
Michael Firstenberg - Assistant Professor of Surgery and Integrative Medicine NorthEast Ohio Medical University, USA & Akron City Hospital - Summa Health System, USA Surgery
\\n\\t
Parul Ichhpujani - MD Government Medical College & Hospital, Department of Ophthalmology, India
Amidou Samie - University of Venda, SA Infectious Diseases
\\n\\t
Shailendra K. Saxena - CSIR-Centre for Cellular and Molecular Biology, Hyderabad, India Infectious Diseases
\\n\\t
Dan T. Simionescu - Department of Bioengineering, Clemson University, Clemson SC, USA Stem Cell Research, Tissue Engineering and Regenerative Medicine
\\n\\t
Ke Xu - Tianjin Lung Cancer Institute Tianjin Medical University General Hospital Tianjin, China Oncology
\\n
\\n\\n
Ophthalmology
\\n\\n
\\n\\t
Hojjat Ahmadzadehfar - University Hospital Bonn Department of Nuclear Medicine Bonn, Germany Medical Diagnostics, Engineering Technology and Telemedicine
\\n\\t
Miroslav Blumenberg - Department of Ronald O. Perelman Department of Dermatology; Department of Biochemistry and Molecular Pharmacology, Dermatology, NYU School of Medicine, NY, USA Dermatology
\\n\\t
Wilfred Bonney - University of Dundee, Scotland, UK Medical Diagnostics, Engineering Technology and Telemedicine
\\n\\t
Christakis Constantinides - Department of Cardiovascular Medicine University of Oxford, Oxford, UK Medical Diagnostics, Engineering Technology and Telemedicine
\\n\\t
Atef Mohamed Mostafa Darwish - Department of Obstetrics and Gynecology , Faculty of Medicine, Assiut University, Egypt Gynecology
\\n\\t
Ana Polona Mivšek - University of Ljubljana, Ljubljana, Slovenia Midwifery
\\n\\t
Gyula Mozsik - First Department of Medicine, Medical and Health Centre, University of Pécs, Hungary
\\n\\t
Shimon Rumelt - Western Galilee-Nahariya Medical Center, Nahariya, Israel Ophthalmology
\\n\\t
Marcelo Saad - S. Paulo Medical College of Acupuncture, SP, Brazil Complementary and Alternative Medicine
\\n\\t
Minoru Tomizawa - National Hospital Organization Shimoshizu Hospital, Japan Gastroenterology
\\n\\t
Pierre Vereecken - Centre Hospitalier Valida and Cliniques Universitaires Saint-Luc, Belgium Dermatology
\\n
\\n\\n
Gastroenterology
\\n\\n
\\n\\t
Hany Aly - Director, Division of Newborn Services The George Washington University Hospital Washington, USA Pediatrics
\\n\\t
Yannis Dionyssiotis - National and Kapodistrian University of Athens, Greece Orthopedics, Rehabilitation and Physical Medicine
\\n\\t
Alina Gonzales- Quevedo Instituto de Neurología y Neurocirugía Havana, Cuba Mental and Behavioural Disorders and Diseases of the Nervous System
\\n\\t
Margarita Guenova - National Specialized Hospital for Active Treatment of Haematological Diseases, Bulgaria
\\n\\t
Eliska Potlukova - Clinic of Medicine, University Hospital Basel, Switzerland Edocrinology
\\n\\t
Raymond L. Rosales -The Royal and Pontifical University of Santo Tomas, Manila, Philippines & Metropolitan Medical Center, Manila, Philippines & St. Luke's Medical Center International Institute in Neuroscience, Quezon City, Philippines Mental and Behavioural Disorders and Diseases of the Nervous System
\\n\\t
Alessandro Rozim - Zorzi University of Campinas, Departamento de Ortopedia e Traumatologia, Campinas, SP, Brazil Orthopedics, Rehabilitation and Physical Medicine
\\n\\t
Dieter Schoepf - University of Bonn, Germany Mental and Behavioural Disorders and Diseases of the Nervous System
\\n
\\n\\n
Hematology
\\n\\n
\\n\\t
Hesham Abd El-Dayem - National Liver Institute, Menoufeyia University, Egypt Hepatology
\\n\\t
Fayez Bahmad - Health Science Faculty of the University of Brasilia Instructor of Otology at Brasilia University Hospital Brasilia, Brazil Otorhinolaryngology
\\n\\t
Peter A. Clark - Saint Joseph's University Philadelphia, Pennsylvania, USA Bioethics
\\n\\t
Celso Pereira - Coimbra University, Coimbra, Portugal Immunology, Allergology and Rheumatology
\\n\\t
Luis Rodrigo - Asturias Central University Hospital (HUCA) School of Medicine, University of Oviedo, Oviedo, Spain Hepatology & Gastroenterology
\\n\\t
Dennis Wat - Liverpool Heart and Chest Hospital NHS Foundation Trust, UK Pulmonology
\\n
\\n\\n
Social Sciences and Humanities Board
\\n\\n
Business, Management and Economics
\\n\\n
\\n\\t
Vito Bobek - University of Applied Sciences, FH Joanneum, Graz, Austria
Joao Luis Garcia Rosa - Associate Professor Bio-inspired Computing Laboratory (BioCom) Department of Computer Science University of Sao Paulo (USP) at Sao Carlos, Brazil
\n\t
Jan Valdman - Institute of Mathematics and Biomathematics, University of South Bohemia, České Budějovice, Czech Republic Institute of Information Theory and Automation of the ASCR, Prague, Czech Republic
\n
\n\n
Earth and Planetary Science
\n\n
\n\t
Jill S. M. Coleman - Department of Geography, Ball State University, Muncie, IN, USA
\n\t
İbrahim Küçük Erciyes - Üniversitesi Department of Astronomy and Space Sciences Melikgazi, Kayseri, Turkey
\n\t
Pasquale Imperatore - Electromagnetic Environmental Sensing (IREA), Italian National Council of Research (CNR), Naples, Italy
\n\t
Mohammad Mokhtari - Director of National Center for Earthquake Prediction International Institute of Earthquake Engineering and Seismology (IIEES), Tehran, Iran
\n
\n\n
Engineering
\n\n
\n\t
Narottam Das - University of Southern Queensland, Australia
\n\t
Jose Ignacio Huertas - Energy and Climate Change Research Group; Instituto Tecnológico y Estudios Superiores de Monterrey, Mexico
Likun Pan - Engineering Research Center for Nanophotonics and Advanced Instrument, Ministry of Education, Department of Physics, East China Normal University, China
\n\t
Mukul Chandra Paul - Central Glass & Ceramic Research Institute Jadavpur, Kolkata, India
\n\t
Stephen E. Saddow - Electrical Engineering Department, University of South Florida, USA
\n\t
Ali Demir Sezer - Marmara University, Faculty of Pharmacy, Department of Pharmaceutical Biotechnology, İstanbul, Turkey
\n\t
Krzysztof Zboinski - Warsaw University of Technology, Faculty of Transport, Warsaw, Poland
\n
\n\n
Materials Science
\n\n
\n\t
Vadim Glebovsky - Senior Researcher, Institute of Solid State Physics, Chernogolovka, Russia Expert of the Russian Fund for Basic Research, Moscow, Russia
\n\t
Jianjun Liu - State Key Laboratory of High Performance Ceramics and Superfine Microstructure of Shanghai Institute of Ceramics, Chinese Academy of Sciences, China
\n\t
Pietro Mandracci - Department of Applied Science and Technology, Politecnico di Torino, Italy
\n\t
Waldemar Alfredo Monteiro - Instituto de Pesquisas Energéticas e Nucleares Materials Science and Technology Center (MSTC) São Paulo, SP, Brazil
Toshio Ogawa - Department of Electrical and Electronic Engineering, Shizuoka Institute of Science and Technology, Toyosawa, Fukuroi, Shizuoka, Japan
\n
\n\n
Mathematics
\n\n
\n\t
Paul Bracken - Department of Mathematics University of Texas, Edinburg, TX, USA
\n
\n\n
Nanotechnology and Nanomaterials
\n\n
\n\t
Muhammad Akhyar - Farrukh Nano-Chemistry Lab. Registrar, GC University Lahore, Pakistan
\n\t
Khan Maaz - Chinese Academy of Sciences, China & The Pakistan Institute of Nuclear Science and Technology, Pakistan
\n
\n\n
Physics
\n\n
\n\t
Izabela Naydenova - Lecturer, School of Physics Principal Investigator, IEO Centre College of Sciences and Health Dublin Institute of Technology Dublin, Ireland
\n\t
Mitsuru Nenoi - National Institute of Radiological Sciences, Japan
\n\t
Christos Volos - Physics Department, Aristotle University of Thessaloniki, Greece
\n
\n\n
Robotics
\n\n
\n\t
Alejandra Barrera - Instituto Tecnológico Autónomo de México, México
\n\t
Dusan M. Stipanovic - Department of Industrial and Enterprise Systems Engineering, University of Illinois at Urbana-Champaign
\n\t
Andrzej Zak - Polish Naval Academy Faculty of Navigation and Naval Weapons Institute of Naval Weapons and Computer Science, Gdynia, Poland
Petr Konvalina - Faculty of Agriculture, University of South Bohemia in České Budějovice, Czech Republic
\n
\n\n
Biochemistry, Genetics and Molecular Biology
\n\n
\n\t
Chunfa Huang - Saint Louis University, Saint Louis, USA
\n\t
Michael Kormann - University Children's Clinic Department of Pediatrics I, Pediatric Infectiology & Immunology, Translational Genomics and Gene Therapy in Pediatrics, University of Tübingen, Tübingen, Germany
\n\t
Bin WU - Ph.D. HCLD Scientific Laboratory Director, Assisted Reproductive Technology Arizona Center for Reproductive Endocrinology and Infertility Tucson, Arizona , USA
\n
\n\n
Environmental Sciences
\n\n
\n\t
Juan A. Blanco - Senior Researcher & Marie Curie Research Fellow Dep. Ciencias del Medio Natural, Universidad Publica de Navarra Campus de Arrosadia, Pamplona, Navarra, Spain
\n\t
Mikkola Heimo - University of Eastern Finland, Kuopio, Finland
\n\t
Bernardo Llamas Moya - Politechnical University of Madrid, Spain
\n\t
Toonika Rinken - Department of Environmental Chemistry, University of Tartu, Estonia
\n
\n\n
Immunology and Microbiology
\n\n
\n\t
Dharumadurai Dhanasekaran - Department of Microbiology, School of Life Sciences, Bharathidasan University, India
Isabel Gigli - Facultad de Agronomia-UNLPam, Argentina
\n\t
Milad Manafi - Department of Animal Science, Faculty of Agricultural Sciences, Malayer University, Malayer, Iran
\n\t
Rita Payan-Carreira - Universidade de Trás-os-Montes e Alto Douro, Departamento de Zootecnia, Portugal
\n
\n\n
Medicine
\n\n
\n\t
Mazen Almasri - King Abdulaziz University, Faculty of Dentistry Jeddah, Saudi Arabia Dentistry
\n\t
Craig Atwood - University of Wisconsin-Madison, USA Stem Cell Research, Tissue Engineering and Regenerative Medicine
\n\t
Oreste Capelli - Clinical Governance, Local Health Authority, Modena, Italy Public Health
\n\t
Michael Firstenberg - Assistant Professor of Surgery and Integrative Medicine NorthEast Ohio Medical University, USA & Akron City Hospital - Summa Health System, USA Surgery
\n\t
Parul Ichhpujani - MD Government Medical College & Hospital, Department of Ophthalmology, India
Amidou Samie - University of Venda, SA Infectious Diseases
\n\t
Shailendra K. Saxena - CSIR-Centre for Cellular and Molecular Biology, Hyderabad, India Infectious Diseases
\n\t
Dan T. Simionescu - Department of Bioengineering, Clemson University, Clemson SC, USA Stem Cell Research, Tissue Engineering and Regenerative Medicine
\n\t
Ke Xu - Tianjin Lung Cancer Institute Tianjin Medical University General Hospital Tianjin, China Oncology
\n
\n\n
Ophthalmology
\n\n
\n\t
Hojjat Ahmadzadehfar - University Hospital Bonn Department of Nuclear Medicine Bonn, Germany Medical Diagnostics, Engineering Technology and Telemedicine
\n\t
Miroslav Blumenberg - Department of Ronald O. Perelman Department of Dermatology; Department of Biochemistry and Molecular Pharmacology, Dermatology, NYU School of Medicine, NY, USA Dermatology
\n\t
Wilfred Bonney - University of Dundee, Scotland, UK Medical Diagnostics, Engineering Technology and Telemedicine
\n\t
Christakis Constantinides - Department of Cardiovascular Medicine University of Oxford, Oxford, UK Medical Diagnostics, Engineering Technology and Telemedicine
\n\t
Atef Mohamed Mostafa Darwish - Department of Obstetrics and Gynecology , Faculty of Medicine, Assiut University, Egypt Gynecology
\n\t
Ana Polona Mivšek - University of Ljubljana, Ljubljana, Slovenia Midwifery
\n\t
Gyula Mozsik - First Department of Medicine, Medical and Health Centre, University of Pécs, Hungary
\n\t
Shimon Rumelt - Western Galilee-Nahariya Medical Center, Nahariya, Israel Ophthalmology
\n\t
Marcelo Saad - S. Paulo Medical College of Acupuncture, SP, Brazil Complementary and Alternative Medicine
\n\t
Minoru Tomizawa - National Hospital Organization Shimoshizu Hospital, Japan Gastroenterology
\n\t
Pierre Vereecken - Centre Hospitalier Valida and Cliniques Universitaires Saint-Luc, Belgium Dermatology
\n
\n\n
Gastroenterology
\n\n
\n\t
Hany Aly - Director, Division of Newborn Services The George Washington University Hospital Washington, USA Pediatrics
\n\t
Yannis Dionyssiotis - National and Kapodistrian University of Athens, Greece Orthopedics, Rehabilitation and Physical Medicine
\n\t
Alina Gonzales- Quevedo Instituto de Neurología y Neurocirugía Havana, Cuba Mental and Behavioural Disorders and Diseases of the Nervous System
\n\t
Margarita Guenova - National Specialized Hospital for Active Treatment of Haematological Diseases, Bulgaria
\n\t
Eliska Potlukova - Clinic of Medicine, University Hospital Basel, Switzerland Edocrinology
\n\t
Raymond L. Rosales -The Royal and Pontifical University of Santo Tomas, Manila, Philippines & Metropolitan Medical Center, Manila, Philippines & St. Luke's Medical Center International Institute in Neuroscience, Quezon City, Philippines Mental and Behavioural Disorders and Diseases of the Nervous System
\n\t
Alessandro Rozim - Zorzi University of Campinas, Departamento de Ortopedia e Traumatologia, Campinas, SP, Brazil Orthopedics, Rehabilitation and Physical Medicine
\n\t
Dieter Schoepf - University of Bonn, Germany Mental and Behavioural Disorders and Diseases of the Nervous System
\n
\n\n
Hematology
\n\n
\n\t
Hesham Abd El-Dayem - National Liver Institute, Menoufeyia University, Egypt Hepatology
\n\t
Fayez Bahmad - Health Science Faculty of the University of Brasilia Instructor of Otology at Brasilia University Hospital Brasilia, Brazil Otorhinolaryngology
\n\t
Peter A. Clark - Saint Joseph's University Philadelphia, Pennsylvania, USA Bioethics
\n\t
Celso Pereira - Coimbra University, Coimbra, Portugal Immunology, Allergology and Rheumatology
\n\t
Luis Rodrigo - Asturias Central University Hospital (HUCA) School of Medicine, University of Oviedo, Oviedo, Spain Hepatology & Gastroenterology
\n\t
Dennis Wat - Liverpool Heart and Chest Hospital NHS Foundation Trust, UK Pulmonology
\n
\n\n
Social Sciences and Humanities Board
\n\n
Business, Management and Economics
\n\n
\n\t
Vito Bobek - University of Applied Sciences, FH Joanneum, Graz, Austria
Denis Erasga - De La Salle University, Phillippines
\n\t
Rosario Laratta - Associate Professor of Social Policy and Development Graduate School of Governance Studies, Meiji University, Japan
\n
\n\n
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I am also a member of the team in charge for the supervision of Ph.D. students in the fields of development of silicon based planar waveguide sensor devices, study of inelastic electron tunnelling in planar tunnelling nanostructures for sensing applications and development of organotellurium(IV) compounds for semiconductor applications. I am a specialist in data analysis techniques and nanosurface structure. I have served as the editor for many books, been a member of the editorial board in science journals, have published many papers and hold many patents.",institutionString:null,institution:{name:"Sheffield Hallam University",country:{name:"United Kingdom"}}},{id:"54525",title:"Prof.",name:"Abdul Latif",middleName:null,surname:"Ahmad",slug:"abdul-latif-ahmad",fullName:"Abdul Latif Ahmad",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"20567",title:"Prof.",name:"Ado",middleName:null,surname:"Jorio",slug:"ado-jorio",fullName:"Ado Jorio",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Universidade Federal de Minas Gerais",country:{name:"Brazil"}}},{id:"47940",title:"Dr.",name:"Alberto",middleName:null,surname:"Mantovani",slug:"alberto-mantovani",fullName:"Alberto Mantovani",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"12392",title:"Mr.",name:"Alex",middleName:null,surname:"Lazinica",slug:"alex-lazinica",fullName:"Alex Lazinica",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/12392/images/7282_n.png",biography:"Alex Lazinica is the founder and CEO of IntechOpen. After obtaining a Master's degree in Mechanical Engineering, he continued his PhD studies in Robotics at the Vienna University of Technology. Here he worked as a robotic researcher with the university's Intelligent Manufacturing Systems Group as well as a guest researcher at various European universities, including the Swiss Federal Institute of Technology Lausanne (EPFL). During this time he published more than 20 scientific papers, gave presentations, served as a reviewer for major robotic journals and conferences and most importantly he co-founded and built the International Journal of Advanced Robotic Systems- world's first Open Access journal in the field of robotics. Starting this journal was a pivotal point in his career, since it was a pathway to founding IntechOpen - Open Access publisher focused on addressing academic researchers needs. Alex is a personification of IntechOpen key values being trusted, open and entrepreneurial. Today his focus is on defining the growth and development strategy for the company.",institutionString:null,institution:{name:"TU Wien",country:{name:"Austria"}}},{id:"19816",title:"Prof.",name:"Alexander",middleName:null,surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/19816/images/1607_n.jpg",biography:"Alexander I. Kokorin: born: 1947, Moscow; DSc., PhD; Principal Research Fellow (Research Professor) of Department of Kinetics and Catalysis, N. Semenov Institute of Chemical Physics, Russian Academy of Sciences, Moscow.\r\nArea of research interests: physical chemistry of complex-organized molecular and nanosized systems, including polymer-metal complexes; the surface of doped oxide semiconductors. He is an expert in structural, absorptive, catalytic and photocatalytic properties, in structural organization and dynamic features of ionic liquids, in magnetic interactions between paramagnetic centers. The author or co-author of 3 books, over 200 articles and reviews in scientific journals and books. He is an actual member of the International EPR/ESR Society, European Society on Quantum Solar Energy Conversion, Moscow House of Scientists, of the Board of Moscow Physical Society.",institutionString:null,institution:{name:"Semenov Institute of Chemical Physics",country:{name:"Russia"}}},{id:"62389",title:"PhD.",name:"Ali Demir",middleName:null,surname:"Sezer",slug:"ali-demir-sezer",fullName:"Ali Demir Sezer",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/62389/images/3413_n.jpg",biography:"Dr. Ali Demir Sezer has a Ph.D. from Pharmaceutical Biotechnology at the Faculty of Pharmacy, University of Marmara (Turkey). He is the member of many Pharmaceutical Associations and acts as a reviewer of scientific journals and European projects under different research areas such as: drug delivery systems, nanotechnology and pharmaceutical biotechnology. Dr. Sezer is the author of many scientific publications in peer-reviewed journals and poster communications. Focus of his research activity is drug delivery, physico-chemical characterization and biological evaluation of biopolymers micro and nanoparticles as modified drug delivery system, and colloidal drug carriers (liposomes, nanoparticles etc.).",institutionString:null,institution:{name:"Marmara University",country:{name:"Turkey"}}},{id:"61051",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"100762",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"St David's Medical Center",country:{name:"United States of America"}}},{id:"107416",title:"Dr.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Texas Cardiac Arrhythmia",country:{name:"United States of America"}}},{id:"64434",title:"Dr.",name:"Angkoon",middleName:null,surname:"Phinyomark",slug:"angkoon-phinyomark",fullName:"Angkoon Phinyomark",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/64434/images/2619_n.jpg",biography:"My name is Angkoon Phinyomark. I received a B.Eng. degree in Computer Engineering with First Class Honors in 2008 from Prince of Songkla University, Songkhla, Thailand, where I received a Ph.D. degree in Electrical Engineering. My research interests are primarily in the area of biomedical signal processing and classification notably EMG (electromyography signal), EOG (electrooculography signal), and EEG (electroencephalography signal), image analysis notably breast cancer analysis and optical coherence tomography, and rehabilitation engineering. I became a student member of IEEE in 2008. During October 2011-March 2012, I had worked at School of Computer Science and Electronic Engineering, University of Essex, Colchester, Essex, United Kingdom. In addition, during a B.Eng. I had been a visiting research student at Faculty of Computer Science, University of Murcia, Murcia, Spain for three months.\n\nI have published over 40 papers during 5 years in refereed journals, books, and conference proceedings in the areas of electro-physiological signals processing and classification, notably EMG and EOG signals, fractal analysis, wavelet analysis, texture analysis, feature extraction and machine learning algorithms, and assistive and rehabilitative devices. I have several computer programming language certificates, i.e. Sun Certified Programmer for the Java 2 Platform 1.4 (SCJP), Microsoft Certified Professional Developer, Web Developer (MCPD), Microsoft Certified Technology Specialist, .NET Framework 2.0 Web (MCTS). 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