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1. Introduction
Groundwater management is an issue which remains a practical matter in many human regions throughout the world [1]. Besides, it is very necessary to clarify that groundwater represents the largest stock of accessible freshwater and accounts for about one-third of freshwater withdrawals globally [2, 3, 4]. However, increased rainfall scarcities have resulted in an augmented use of groundwater, in order to satisfy the increasing domestic, agricultural, and environmental-ecosystem preservation for different water.
Nevertheless, it is necessary to take into account that historically, surface water has been the main source of water for human consumption, as it was easy and cost effective to access. So, it can be expected that during the second half of the twentieth century, groundwater withdrawals will increase. It is also very relevant to reflect that groundwater supply could represent around one third of the world population [5].
This wide use of groundwater in many parts of the world has resulted in water level decline and groundwater depletion and is mainly related to phenomena such as biodiversity loss, pollution, and seawater intrusion in coastal aquifers. An example could be found in the paper by El Moujabber et al. [6], in which the state of groundwater desalination by seawater intrusion in the Lebanese cost is introduced (specifically in the region of Choueifat-Rmeyle, located in the south of Mount-Lebanon). The main consequence that is obtained is related to the fact that groundwater management can behave like relevant backstop technologies and also that substitutes have become a practical concern in many arid and semiarid regions throughout the world [7].
A fundamental idea that needs to be pointed out is that groundwater is essential for sustaining agriculture production patterns, as well as consumption models and the biodiversity or the resilience of ecosystems. The combination of this fact with the intense scarcity in many parts of the world makes necessary the development of rules for the corrected and efficient allocation of resources among competing uses over time and space.
This presents an economic question which has been close to groundwater economics since the middle years of the decade of 1950s. It is necessary to point out that the question of how to manage this resource, mainly because groundwater constitutes about 89% of the freshwater on earth (discounting that in the polar ice caps). From this, an important economic concept could be deduced related to water scarcity and which is related to the fact that the world water scarcity is one of the most important hydraulic resources that need to be taken into account.
It is also necessary to point out that groundwater systems are rather dynamic with groundwater in motion from zones of recharge to areas of discharge and that a great number of years could, hundreds of years, interfere in the passage of water through this subterranean part of the hydrological cycle. Since flow rates regularly do not ordinarily go beyond a small number of meters per day and can be as low as 1 meter per year (these groundwater velocities compare to rates of up to 1 meter per second for river flows) [8].
Groundwater resources provide a primary (or supplemental) source of irrigation water throughout much of the world, yet overpumping and subsequent aquifer depletion may pose “the single largest threat to irrigated agriculture” [9, 10].
Two main questions need to be indicated in when taking water extractions into account. The first is double: one is water scarcity in local watersheds (or whole basins created by extreme surface and groundwater withdrawals). The other is water degradation from pollution loads leading to many tracts of rivers and whole aquifers being damaged and losing their capacity to sustain ecosystem functioning and human accomplishments.
Following the wide-scale development of groundwater pumping for agriculture in 1950s, some results have been obtained that the open access nature of groundwater implied that farmers were overextracting water, and therefore, it could be exhausted much before than it might be economically optimal.
These conclusions were called into question by Gisser and Sanchez. These authors, mainly in their very influential paper argue that the difference in producer surplus between the open access and optimally managed cases was numerically insignificant for large aquifers subject to inelastic water demand. Perhaps the most interesting point in the work by Gisser and Sánchez is multidisciplinarity. An essential assumption that we need to take into account is that the GSE model is a dynamic model. Besides, we need to take into account that variables in the model are economic, hydrological, and agronomic variables of groundwater use. In this chapter, the demand and supply functions for irrigated water are defined, and these functions are associated with the hydrological characteristics of the aquifer. Then, the path of water allocation through time is calculated under the policy regime and the free-market regime [11].
This effect has remained controversial, and numerous studies have analyzed whether the Gisser and Sanchez Effect (GSE) persists under a variety of specific conditions, such as convex pumping costs [12], shifting (nonconstant) water demand [13], adaptation by crop shifting [14], confined aquifers [15], heterogeneous users [16]; strategic decision-making [17, 18], conjunctive management [19], risk aversion [20], and backstop water sources [21]. These studies generally find support for the GSE, even under all these different conditions.
Nevertheless, it is necessary to point out that authors, such as Stratton et al. [22], apply the GSE model, although relaxing a very important significant assumption of a fixed irrigation technology. Results indicate that the GSE fails when irrigation technologies with different water use efficiency become available. These results are robust and hold even when maintaining some of the very fundamental statements in the original model (such as constant marginal pumping costs per linear foot of lift). Besides, the gains from optimal groundwater management become even more significant when irrigation technology is not only variable but also endogenous variables. That is, variables whose values the model is designed to explain. In the model, there are also exogenous variables. That is, variables whose values are taken as given from outside the model [23]. The expression “Endogenous Technical Change” implies that higher water costs could induce the development of technologies that might improve water use efficiency [19, 24]. The expression “Endogenous Technical Change” implies that higher water costs could encourage the development of technologies that might improve water use efficiency [24, 25].
The main objective of this chapter is to re-evaluate the validity of the GSE hypothesis in groundwater management. In this chapter, the conceptual framework within which the elements interacting in the management of groundwater resources is examined. The most important conclusion obtained is that the role of the market is limited with respect to the price of water in an aquifer. This is an important result, because it points to the mechanism that could pull competitive water prices and quality-graded quantity of groundwater, in line with their equilibrium levels. In Section 2, some models of groundwater use and management are introduced, and the most important economic models for groundwater use can be found (joint with the potential of groundwater management control variables in such models). In Chapter 3, some relationships between the Gisser and Sanchez effect and the difficulties to establish clear groundwater property rights are discussed. In Section 4, the robustness of GSE under a private if property rights regime is discussed, both in quantity and in quality terms. In Section 5, a discussion section is introduced. Finally, some conclusions are provided.
2. Some models of groundwater use and the potential for groundwater management
It is necessary to take into account that implicit in the different concerns about groundwater, an essential principle can be found. This is related to the fact that if no intervention exists, then groundwater pumping will be mismanaged. Another important point that needs to be pointed out is that if groundwater pumping is inefficient, then, the lack of central (and optimal), control, underlines that the estimates of the welfare loss (under the common property regime) should depend on the specific model of firm behavior which might be enlisted in the analysis. This should allow to conclude in favor of an existing potential and pressing need for the development and implementation of management policies for groundwater resources [32].
It is also interesting to point out that when groundwater withdrawals exceed recharge, water will be mined over time until either supplies are exhausted or the marginal cost of pumping additional water should become extremely expensive [33]. An essential issue related to this assumption is that a marginal user cost is associated with mining groundwater, and this is related to the opportunity cost which is connected with the unavailability in the future of any unit of water used in the present.
A well-organized distribution should consider this user cost, which effectively signals the scarcity of the resource and is called the resource’s scarcity rents. Therefore, efficient pricing of a resource that exhibits natural supply constraints incorporates both marginal cost of extraction and scarcity rents. Scarcity rents must be imposed on current users.
Given the complexity of establishing clear groundwater property rights, scarcity rents are frequently difficult to be recognized and are not easy to be estimated. Some authors in which a discussion about this point could be found are, for instance [31, 34, 35, 36, 37].
Ignoring scarcity rents implies that the price of groundwater is usually too low and extraction is above the socially optimal level. If an optimal dynamic management of common-pool groundwater resources is not considered, or in the presence of a competitive extraction regime ignoring scarcity rents, results in inefficient pricing and misallocation of resources. This essential argument has to do with the way markets behave, and it could perfectly be competitive. Under these circumstances, the problem is not so much with the market mechanism but with the way property rights behave.
3. Groundwater property rights
Given the difficulty of establishing clear groundwater property rights, scarcity rents are frequently difficult to be estimated. Ignoring scarcity rents should imply that groundwater prices could be too low and extraction might be above the socially optimal level. From this, the main conclusion is that, in the absence of optimal dynamic management of common pool groundwater resources, or, alternatively, in the presence of a competitive extraction regime, ignoring scarcity rents, could result in inefficient pricing and misallocation of resources.
Just in the case, there is no optimal dynamic management of common pool groundwater resources, or, alternatively, in the presence of a competitive extraction regime, ignoring scarcity rents results in inefficient pricing competitive extraction regime, inefficient pricing, and misallocation of the resource.
From this, an interesting question might be pointed out: How could be explained that a competitive dynamic solution of groundwater exploitation is almost identical (in terms of derived social welfare) to the efficient management solution, in the way it is claimed by the GSE effect?
3.1 The Gisser-Sánchez effect
The GSE explains a contradictory empirical result, present and persisting in the dynamic solutions of groundwater exploitation under different extraction regimes (since 1980) [1]. In spite of the fact that depletion of aquifers is a major threat to many freshwater ecosystems all over the world, the social benefits from managing groundwater are numerically insignificant. It needs to be pointed out that GSE encompasses to a general rule, and then the role and scope of water management are severely limited. It is also essential to point out that, even if implementing optimal extraction is not going to be costless. In this section, a review of [38] is introduced about the theoretical and empirical attempts to address the GSE and discuss the potential for groundwater management.
3.2 The Gisser-Sanchez model and groundwater management
Problems of groundwater allocation have been studied basically in the context of the theory of mine [26, 27, 28, 29]. The basic model by Gisser and Sanchez is a simplified representation of the economic, hydrologic, and agronomic facts that must be considered relative to the irrigator’s choice of water pumping [1]. The validity of the GSE model rests on the key assumption that the aquifer has to be quite large and on the secondary assumption of a small slope in the water-demand function.
A separate literature should also have to be taken into account, which deals with groundwater quality. Some papers in this line can be found such as [30, 31, 32].
Groundwater allocation problems have been studied mainly in the context of mine and economists like [33, 34, 35]. Some principles of inventory management to derive decision rules for the optimal temporal allocation in a dynamic programming format can also be found in such papers. The effects of different policy instruments that could correct misallocation of commonly owned groundwater can be found in papers such as [31, 35, 36, 37, 38, 39], which studied the effects of different policy instruments that might correct the misallocation of commonly owned groundwater. One of the main results of this chapter is that net benefits from groundwater management could amount to over $100 per acre, but noted that these benefits could decline with increases in interest rate. One of the solutions to this problem was obtained by authors such as Allen and Gisser [40], who derived a formula for a tax that should be imposed on groundwater which was pumped in order to yield the optimal control solution. Finally, in papers such as [41], it can be recognized the issue of congestion externality in aquifers with open access characteristics and suggested a charging tax to accommodate this externality.
When this point is achieved, farmers will either import supplemental water or be restricted to use a smaller amount of water by being assigned water rights. Nevertheless, some changes in the hypothesis related to regulation of water pumping in the aquifer could be made. This case allows to model consistently an optimal control problem and also allows one kind of clarification that should be related with the case of no control. This is the departure point for the works [9, 10] by Gisser and Sanchez.
The basic model analyzed by Gisser and Sanchez is a simplified representation of the economic, hydrologic, and agronomic facts that should be considered for the irrigator’s choice of water pumping. An irrigator benefit function could be represented using this function suggested by [44]:
πt=Vwt−CHtwtE1
where π(t) denotes profits at time t. Net farm revenue from water use π(t) neglecting pumping costs is denoted by
Vw=∫0wpxdxE2
where p(x) is the inverse demand function for water. C(H) is the average and marginal pumping costs per acre-foot of water and H(t) is the height of water table above some arbitrary reference point at time t [1, 40]. The change in the height of water is given by differential Eq. (2), which represents the hydrologic state of the aquifer (or equivalently, the environmental constraint of the problem)
H´=1ASR+a−1w,H0=H0E3
In this equation, R exemplifies a constant recharge determined in acre feet per year; a is the constant return flow coefficient (which could be considered to be just a simple number); H0 is the initial level of the water table measured in feet above sea level; A is the surface area of the aquifer (uniform at all depths), measured in acres per year; and S is the specific yield of the aquifer. These equations are based on the UNESCO-Encyclopedia Life Support Systems and also on the papers by [1, 43, 45] on the Gisser-Sanchez effect.
More precisely, the aquifer in Gisser and Sanchez’s work is modeled as a bathtub, unconfined aquifer, with infinite hydraulic conductivity. It is necessary to point out that infinite hydraulic conductivity implies that the aquifer will never dry up, irrespective of groundwater extraction rates, which is equivalent to the assumption of a bottomless aquifer. The adoption of this hypothesis can be acknowledged by the hypothesis that it is implied by an standard hypothesis which is related to the literature and which implies that time goes to infinity [1]. Nevertheless, if this is not this way, a steady-state solution might not be reached. Besides, Provencher [43] showed that the optimal pumping rate can be substantially lower when the hydraulic conductivity is small enough to result in a significant cone of depression around the well. The assumption of constant return flow in the presence of fixed irrigation technology suggests a constant rate of water application.
The hypothesis of deterministic and constant recharge in conjunction with the hypothesis of constant return flow suggests constant types of land use [44], independence of surface water and groundwater systems, and constant average rainfall. Besides, sunk costs, replacement costs, and capital costs in general are overlooked, and it is implicitly assumed that energy costs are constant. It is also indirectly accepted that the well pump capacity constraint is nonbinding. Finally, refinement in Gisser and Sanchez’s model could be also achieved by assuming that only land superimposing the aquifer can be irrigated. That is, the demand curve does not shift to the right over time. This implies that, the unambiguous recognition of the fact that the main hypothesis behind the GSE indicates that the result should be carefully when working on real aquifer systems.
Given the above hydroeconomic model, Gisser and Sanchez used a linear water demand function (estimated by [31, 32]) using parametric linear programming, hydrologic parameters that were considered realistic in the 1960s, and a discount rate of 10%, and simulated the intertemporal water pumpage for Pecos Basin in New Mexico, once under the assumption of no control and once under the assumption of optimal control. The most interesting result is that the trajectories under the two regimes are almost identical. This result leads to the main conclusion that there is no substantive quantitative difference between socially optimal rules for pumping water and competitive rates. Therefore, the welfare loss from intertemporal misallocation of pumping effort is negligible. This conclusion amounts to the GSE.
An important effect to consider is that, solving analytically the model, Gisser and Sanchez main result is that, if Eq. (3) is true, then the difference between the two strategies is so small that it can be ignored for practical consideration, where Eq. (3) is
kCta−1AS2≃0E4
In Eq. (4), k can be considered to be the reduction in demand for water per $1 intensification in price (that is, the slope of the uncompensated demand curve for groundwater), Ct is the intensification in pumping cost per acre-foot per 1-foot decline in the water table, and AS are given in Eq. (2). If Eq. (3) holds, then the rate of discount will be practically identical with the exponent of the competition result. Therefore, as long as the slope of the groundwater demand is small relative to the aquifer’s area times its storativity [1], GSE will persist. From this, the main conclusion is that, if differences between optimal and competitive rates of water pumping are small, then policy considerations can be limited to those which ensure that the market operates in a competitive fashion, and concerns relative to rectifying common property effects could be removed.
3.3 Robustness of the GSE effect
The GSE effect presents important policy implications. Some empirical papers discussing the robustness of this effect are, Noel et al. [35] found that control increases the value of groundwater in the Yolo basin in California, by 10%. This result is fairly different from [37], who found that control raised the net benefit of groundwater in the Ogallala basin by only 0.3% empirical estimates of benefits from groundwater management in Kern county (California, USA) do not exceed 10%. Nevertheless, in works such as [39], it can be found that groundwater management in the Texas High Plains would be unwarranted, and he proceeded with a sensitivity analysis of present value profits using different slopes and intercept values for the groundwater-demand curve. It is interesting to point out that this analysis indicated that benefits from groundwater management do not increase monotonically as the absolute value of the slope increases.
A basic hypothesis of the Gisser and Sánchez model is that the demand curve for water is linear. This is a fairly conventional hypothesis in most economic demand models. In order to study the relative importance of this hypothesis for the GSE, optimal control and no-control strategies are compared, using a nonlinear demand curve [40]. This comparison confirmed that, for the case of the nonlinear demand function, what had been demonstrated by the GSE for the case of a linear demand function.
However, in works such as [20], it can be found that the differences between the two regimes may not be trivial if the relationship the average extraction cost and the water table level and/or if there exist significant differences in land productivity, applying dynamic programming to a model of a confined aquifer underlying the Crow Creek Valley in South-Western Montana.
It is essential to take into account that when land is assumed to be homogeneous, the gross returns function with respect to water use tends to be nearly linear. Nevertheless, with greater heterogeneity in productivity, the returns function is more concave, and differences in the optimal use policy under a common property setting are more pronounced [1]. Hence, the need for more theoretical work is to determine an asymmetric groundwater pumping differential game, where differences in land productivity are taken into account.
3.4 Variable relations and endogenous rates of change
Implicit in GSE model is the hypothesis of nonvariable economic relations (that is, time-independent demand) and/or exogenous and constant rates of change (that is, constant and fixed exogenous crop mix, constant crop requirements, fixed irrigation technology), and some significant exceptions can be found such as [43, 44], with constant exogenous kinds of land use and nonvariable hydrologic conditions.
Nevertheless, in studies with a long run perspective, predictable results could turn out to be weaker as the steady state is approached. Estimated benefit and cost functions used in the simulations of GSE may bear little relation to the actual benefit and cost functions when economic, hydrologic, and agronomic conditions are much different. More complex representations of increasing resource scarcity incorporate opportunities for adaptation to the rising resource prices which are a main indicator for scarcity. In the long run, adoption of new techniques, substitution of alternative inputs, and production of a different mix of products offer rational responses to increasing scarcity [1], [38].
4. The robustness of GSE under a private property rights regime
The solution which is commonly proposed for the inefficiencies arising in common property resource extraction is central-optimal control by a regulator, who uses taxes or quotas to obtain the efficient allocation of resources over time.
In the background of groundwater depletion, a solution has been commonly suggested which is based on a tradable permit scheme [37, 38]. In the framework of groundwater reduction, a number of authors have recommended a similar institutional arrangement in which firms are arranged and endowment of tradable permits to the in situ groundwater stock, which they control over time. Each firm’s bundle of permits represents its private stock of groundwater.
This private stock is worsening due to groundwater pumping and intensifications to reflect the firm share of periodic recharge. It also changes in response to the activity of the firm in the market for groundwater stock permits, increasing when permits are purchased and decreasing when permits are sold. The market price for permits serves to allocate groundwater over time.
It is necessary to point out that this particular regime is inefficient, mainly because both the pumping cost externality and the risk externality persist after the allocation of permits. Moreover, this regime is time inconsistent. However, different efforts to quantify the value of groundwater resource under both optimal control and the private property rights regime indicate that groundwater privatization recovers most of the potential gain from management. In particular, a programming model for Madera County, in California (USA), can be found in [37]. This regime recovered 95% of the potential gain from management.
4.1 The GSE in models of conjunctive use of surface and groundwater
A tributary aquifer is characterized by a groundwater stock that is hydrologically connected to a body of surface water. In this aquifer, surface water may recharge the underground aquifer, or groundwater may supplement surface flows depending upon hydrological conditions.
In papers such as [38], results can be found in which an analytical economic model is developed and is focused primarily on the hydrologic link between surface and groundwater, by modeling the instantaneous rate of aquifer recharge caused by groundwater pumping, through river effects. In this chapter, some externalities river effects can be found, which reinforced groundwater overpumping present due to the usual common property effects. Results of this chapter indicate that optimal policy requires compensation to be paid for both river effects and aquifer depletion net of river effects. This work points to an externality created by groundwater overpumping provoked mainly by the common property effects.
From this, the main conclusion which needs to be pointed out is that optimal policy requires a recompense to be paid for both river effects and aquifer depletion net of river effects [39]. It is necessary to highlight that these effects indicate the existence of some externalities which could be related to groundwater pumping, which might be adjusted with the precise management. The main consequence probably could be that GSE might be very likely removed by the improvement in management benefits.
Unfortunately, no empirical results exist of these results focusing primarily on the hydrologic link between ground and surface water, and at the same time acknowledging the stochastic nature of surface water supplies. Instead, the main literature that incorporates stochastic surface supplies into a groundwater model in which surface water and groundwater are modeled as substitute goods, aquifers are not connected with surface water, and they only benefit from substantial natural recharge.
5. Discussions
Regarding the GSE model, it needs a number of important assumptions. One of the most significant has to do with the disregard for aquatic ecosystems linked and dependent on aquifer systems.
In the GSE model, a very special point needs to be pointed out, which is that the aquifer is presented as a “bath-tub”, unconfined aquifer, with infinite hydraulic conductivity [40]. A bath-tub approach to modeling an aquifer assumes that it responds uniformly and instantly to groundwater extraction [41, 46, 47]. From this, the spatial distribution of the users of the resource is not so relevant, and the evolution of the spatial profile of drawdown does not affect current and future extraction choices. Gisser-Sánchez assumes a deterministic and constant recharge, constant return flow and average rainfall, independence of surface water and groundwater systems, and a bottom-less aquifer. Since their competitive steady state presents a positive water stock, their estimation of welfare gains from optimal management excludes stock externality [42].
Another important assumption which is discussed in this chapter is the appropriateness of the stock effect assumption. This hypothesis reflects the dependence of extraction costs and the eventual benefits on the stock of the resource. From this, it could be established the way these assumptions might affect the time variation of the shadow price of groundwater externality. In this chapter, the main result is that this could lead to a declining value of in situ resource over time. Therefore, the addition on nonmarginal extraction costs could be close to inappreciable, which could imply the validation of the Gisser-Sanchez effect, which also presents the remarkable hypothesis that groundwater markets have the benefit of allowing more flexible movement of water to serve changing conditions and demands.
Finally, a main conclusion could be derived from the paper introduced which is that a very relevant model such as [4, 5] is a very appropriate work to analyze groundwater management, mainly because it states the conditions under which welfare improvements from policy interventions could be significant in aquifer administration. This result could be compared with nonregulation or free market solutions in groundwater management.
6. Conclusion
The main conclusion of this chapter has to do with the GSE effect and points mainly to the different effects related to welfare improvements and aquifer management. In this work, an optimal policy requires a compensation to be paid for both river effects and aquifer depletion, which points to an additional externality created by groundwater pumping. This externality could be corrected with an appropriate management of the groundwater, which could eventually eliminate the GSE effect and even increase management benefits.
No empirical results have been obtained in order to test these results, which have to do mainly with the eventual links between ground and surface water. These results could be pertinent in order to improve groundwater management, because from this, the stochastic nature of surface water flows could be acknowledged.
Nevertheless, probably the most significant result in this chapter is that different effects related to welfare improvements and aquifer management and the relevance of the GSE effect exists. Besides, it is necessary to indicate that an optimal aquifer management policy requires a compensation to be paid for both the existing river effects and aquifer depletion. These conclusions stem from the fact that externalities exist, which are linked to groundwater pumping. This externality could also be corrected with a suitable management of the groundwater. This result is quite relevant because it could potentially remove the GSE effect, and therefore, even increase management benefits.
It is similarly essential to take into account the appropriateness of some of the assumptions in the model, since some of them (like the linear relationship between pumping costs for nonconsumptive benefits), and which are an essential tool in groundwater management.
Environmental uses of groundwater water and the way markets work present a significant impact on users and the environment. An interesting conclusion is provided by [48], in his paper for the journal Resources (from Resources for the Future), which is that there is rapid depletion of aquifers in the United States, and this presents significant impacts on users and the environment, requiring stakeholders across the country to look for creative and effective policy solutions. So, there is an interesting conclusion that groundwater markets can be applied broadly in groundwater management in order to protect one the most relevant freshwater environmental resource.
\n',keywords:"Gisser-Sanchez effect, groundwater, property rights",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/66379.pdf",chapterXML:"https://mts.intechopen.com/source/xml/66379.xml",downloadPdfUrl:"/chapter/pdf-download/66379",previewPdfUrl:"/chapter/pdf-preview/66379",totalDownloads:458,totalViews:96,totalCrossrefCites:0,totalDimensionsCites:1,hasAltmetrics:0,dateSubmitted:"August 27th 2018",dateReviewed:"February 27th 2019",datePrePublished:"March 26th 2019",datePublished:"May 22nd 2019",dateFinished:"March 26th 2019",readingETA:"0",abstract:"The main subject of this chapter is related to the relevance of the Gisser-Sanchez effect in groundwater. It is important to point out that groundwater resources provide a primary source of irrigation water throughout much of the world. Two main questions need to be indicated when taking water extractions into account. The first has to do with water scarcity in local watersheds or whole basins created by excessive surface and groundwater withdrawals. The other is related to water degradation and the pollution loads leading to many tracts of rivers and whole aquifers being spoiled and losing their capacity to sustain ecosystem functioning and human activities. These conclusions were called into question by the Gisser and Sanchez analysis. These authors argue that the difference in producer surplus between the open access and optimally managed cases was numerically insignificant for large aquifers subject to inelastic water demand. Perhaps the most interesting point in the work by Gisser and Sanchez is multidisciplinarity.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/66379",risUrl:"/chapter/ris/66379",book:{slug:"groundwater-resource-characterisation-and-management-aspects"},signatures:"Oscar Alfranca",authors:null,sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Some models of groundwater use and the potential for groundwater management",level:"1"},{id:"sec_3",title:"3. Groundwater property rights",level:"1"},{id:"sec_3_2",title:"3.1 The Gisser-Sánchez effect",level:"2"},{id:"sec_4_2",title:"3.2 The Gisser-Sanchez model and groundwater management",level:"2"},{id:"sec_5_2",title:"3.3 Robustness of the GSE effect",level:"2"},{id:"sec_6_2",title:"3.4 Variable relations and endogenous rates of change",level:"2"},{id:"sec_8",title:"4. The robustness of GSE under a private property rights regime",level:"1"},{id:"sec_8_2",title:"4.1 The GSE in models of conjunctive use of surface and groundwater",level:"2"},{id:"sec_10",title:"5. Discussions",level:"1"},{id:"sec_11",title:"6. Conclusion",level:"1"}],chapterReferences:[{id:"B1",body:'Koundouri P. Current issues in the economics of groundwater resource world management. Journal of Economic Surveys. 2004;18(5):703-740'},{id:"B2",body:'Gorelick SM, Zheng C. 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Costs of resource depletion externalities: A study of groundwater overexplotation in Andhra Pradesh, India. Environment and Development Economics. 2005;10:533-556'},{id:"B45",body:'Tietenberg T, Lewis L. Environmental Economics & Policy. 10th ed. London: Pearson; 2018'},{id:"B46",body:'Katic PG. Three essays on the economics of groundwater extraction [Ph.D. thesis]. Australia: The Australian National University; 2011'},{id:"B47",body:'Katic PG, Grafton RQ. Economic and spatial modelling of groundwater extraction. Journal of Hydrology. 2012;20:831-834'},{id:"B48",body:'Kuwayama Y. Groundwater markets: Managing a critical, hidden resource, resources. Resources for the Future. 2014;43:1-11'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Oscar Alfranca",address:"oscar.alfranca@upc.edu",affiliation:'
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Differing in Their Capacity to Withstand Salinity",slug:"antioxidant-enzyme-activities-as-a-tool-to-discriminate-ecotypes-of-crithmum-maritimum-l-differing-i",signatures:"Ben Hamed Karim, Magné Christian and Abdelly Chedly",authors:[{id:"87734",title:"Prof.",name:"Chedly",middleName:null,surname:"Abdelly",fullName:"Chedly Abdelly",slug:"chedly-abdelly"}]},{id:"26978",title:"Use of Finite Element Method to Determine the Influence of Land Vehicles Traffic on Artificial Soil Compaction",slug:"use-of-finite-element-method-to-determine-the-influence-of-land-vehicles-traffic-on-artificial-soil-",signatures:"Biris Sorin-Stefan and Vladut Valentin",authors:[{id:"81249",title:"Dr.",name:"Sorin-Stefan",middleName:"I",surname:"Biris",fullName:"Sorin-Stefan Biris",slug:"sorin-stefan-biris"},{id:"149766",title:"Dr.",name:"Valentin",middleName:null,surname:"Vladut",fullName:"Valentin Vladut",slug:"valentin-vladut"}]},{id:"26979",title:"The Influence of Water Stress on Yield and Related Characteristics in Inbred Quality Protein Maize Lines and Their Hybrid Progeny",slug:"the-influence-of-water-stress-on-yield-and-related-characteristics-in-inbred-quality-protein-maize-l",signatures:"Dagne Wegary, Maryke Labuschagne and Bindiganavile Vivek",authors:[{id:"81395",title:"Prof.",name:"Maryke",middleName:"Tine",surname:"Labuschagne",fullName:"Maryke Labuschagne",slug:"maryke-labuschagne"}]},{id:"26980",title:"Application of Molecular Breeding for Development of the Drought-Tolerant Genotypes in Wheat",slug:"application-of-molecular-breeding-for-development-of-the-drought-tolerant-genotypes-in-wheat-",signatures:"Mohamed Najeb Barakat and Abdullah Abdullaziz Al-Doss",authors:[{id:"85469",title:"Prof.",name:"Mohamed Najib",middleName:null,surname:"Barakat",fullName:"Mohamed Najib Barakat",slug:"mohamed-najib-barakat"},{id:"124809",title:"Prof.",name:"Abdullah Abdullaziz",middleName:null,surname:"Al-Doss",fullName:"Abdullah Abdullaziz Al-Doss",slug:"abdullah-abdullaziz-al-doss"}]},{id:"26981",title:"Integrated Agronomic Crop Managements to Improve Tef Productivity Under Terminal Drought",slug:"integrated-agronomic-crop-managments-to-improve-tef-productivity-under-terminal-drought",signatures:"Dejene K. Mengistu and Lemlem S. Mekonnen",authors:[{id:"83613",title:"Mr.",name:"Dejene K.",middleName:"Kassahun",surname:"Mengistu",fullName:"Dejene K. Mengistu",slug:"dejene-k.-mengistu"},{id:"119640",title:"Mrs.",name:"Lemlem S.",middleName:null,surname:"Mekonnen",fullName:"Lemlem S. Mekonnen",slug:"lemlem-s.-mekonnen"}]},{id:"26982",title:"Sugarcane Responses at Water Deficit Conditions",slug:"sugarcane-responses-at-water-deficit-conditions-",signatures:"Sonia Marli Zingaretti, Fabiana Aparecida Rodrigues, José Perez da Graça, Livia de Matos Pereira and Mirian Vergínia Lourenço",authors:[{id:"85115",title:"Prof.",name:"Sonia Marli",middleName:null,surname:"Zingaretti",fullName:"Sonia Marli Zingaretti",slug:"sonia-marli-zingaretti"},{id:"124439",title:"Dr.",name:"Mirian Vergínia",middleName:null,surname:"Lourenço",fullName:"Mirian Vergínia Lourenço",slug:"mirian-verginia-lourenco"},{id:"124440",title:"Dr.",name:"Fabiana Aparecida",middleName:null,surname:"Rodrigues",fullName:"Fabiana Aparecida Rodrigues",slug:"fabiana-aparecida-rodrigues"},{id:"124441",title:"Dr.",name:"Livia",middleName:null,surname:"De Matos Pereira",fullName:"Livia De Matos Pereira",slug:"livia-de-matos-pereira"},{id:"124442",title:"MSc.",name:"Jose",middleName:null,surname:"Perez da Graça",fullName:"Jose Perez da Graça",slug:"jose-perez-da-graca"}]},{id:"26983",title:"Strategies for Selecting Drought Tolerant Germplasm in Forage Legume Species",slug:"strategies-for-selecting-drought-tolerant-germplasm-in-forage-legume-species",signatures:"Hernán Acuña, Luis Inostroza and Gerardo Tapia",authors:[{id:"84028",title:"Dr.",name:"Hernán",middleName:null,surname:"Acuña",fullName:"Hernán Acuña",slug:"hernan-acuna"},{id:"149813",title:"Dr.",name:"Luis",middleName:null,surname:"Inostroza",fullName:"Luis Inostroza",slug:"luis-inostroza"},{id:"149814",title:"Dr.",name:"Gerardo",middleName:null,surname:"Tapia",fullName:"Gerardo Tapia",slug:"gerardo-tapia"}]}]}]},onlineFirst:{chapter:{type:"chapter",id:"74600",title:"Blister Blight Disease of Tea: An Enigma",doi:"10.5772/intechopen.95362",slug:"blister-blight-disease-of-tea-an-enigma",body:'\n
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1. Introduction
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Tea is one of the most popular beverages worldwide, having gained popularity for its taste, stimulating effect, various medicinal properties, and related health benefits. Tea is processed from the leaves of evergreen, woody, and perennial tea plants (Camellia sinensis) belonging to the family Theaceae. Three indigenous varieties of tea plant are found viz. C. sinensis (L.) O. Kuntze (China type), C. assamica (Assam type), and (3) C. assamica sub spp lasiocalyx (Planchon ex Watt.) or Cambod type. These varieties are capable of cross-pollination and interbreeding, resulting in heterogenous hybrids. Under natural conditions, the tea plants can grow up to a maximum height of 15 m, while cultivated tea plantations are maintained as a bush with a height of 60–100 cm, which facilitates the plucking of tender leaves [1]. Tea was first used as a beverage in China in 2737 B.C and was introduced in India for commercial production by the erstwhile colonial British, through the East India Company in 1853 [2]. Tea plantations in India are found in three main geographic regions - the Northeast, (Assam, West Bengal, Tripura, Sikkim, Manipur, Nagaland, Meghalaya, Arunachal Pradesh, and Mizoram), the South (Karnataka and Tamil Nadu), and the Northwest (Himachal Pradesh and Uttarakhand) [3]. The Darjeeling Tea from northeast India has been signified as the world’s premium, and exotically flavored tea owing to its unique flavor and aroma, earning itself a GI tag.
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Presently tea is cultivated worldwide across 61 countries of which China, India, Kenya, Sri Lanka, and Vietnam are the largest tea-producing countries, contributing 77% of world production and 80% of global exports (Figures 1 and 2). China is reported to produce 2700 million kg of tea of which 366.6 million Kg were exported for the year 2019 with one million hectares under tea cultivation. This was followed by India and Kenya with 1390.1 and 458.9 million kg of tea production. Recently, Kenya was listed with the highest exporter of tea for the year 2019 with 392.6 million kg of exports. India accounts for 23% of the total world tea production with an area of 400,000 hectares under tea cultivation. This contributes to about US $803 million to the Indian economy (Tea board of India). India is followed by Sri Lanka with an export worth US $721.3 million. In addition to contributing majorly to the economies of the tea growing countries, the tea industry also provides livelihood to a significant part of the population in these countries.
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Figure 1.
Tea producing countries worldwide (a), area of tea cultivation in India (b).
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Figure 2.
Tea production of top 10 countries for the year 2019–2020 (source: Tea board of India).
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Different forms of tea have been produced from the same tea plantations depending on the method of processing and plucking of leaves. Some of them are black tea, green tea, oolong tea, white tea, pure tea, and dark tea. Black tea is the most popular form of tea produced in all the major tea producing countries including India, Kenya, and Sri Lanka. Production of black tea in India accounts for 85% of total worldwide production and green tea is being produced by a few tea gardens. Green tea is the most popular form of tea in China followed by black tea.
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Tea being a perennial and monoculture crop, the microclimate of tea plantations makes it prone to various pests and pathogens [4]. Chen and Chen recorded around 400 pathogens 507 species of fungi infecting tea plants [5, 6]. Although all the parts of the tea plant including leaf, stem, and root are prone to infection, the pathogens invading leaf parts are of great concern as the main source of commercial production of tea is the young and fresh leaves. The incidence of diseases in the leaves significantly affects the crop yield and quality of made tea. This also directly affects the economy of agronomic countries where tea is considered an important cash crop. The various diseases of tea can be categorized into primary and secondary diseases. In case of the primary diseases, the pathogens directly invade healthy tea bushes while secondary diseases are caused by weaker parasites infecting already diseased/infected tea bushes. In this context, some of the most important diseases infecting the leaf, stem, and root of tea plantations are listed in Table 1.
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Blister blight disease is one of the most serious primary foliar tea diseases that significantly affects the crop yield and quality throughout various regions of tea-producing countries across the world. The causal organism of Blister blight disease is the biotrophic fungus Exobasidium vexans Massee. Peal was the first to recognize the occurrence of blister blight in the year 1868 in North East India [9]. The disease mainly attacks young harvestable tender leaves which are used for the commercial production of tea. Blister blight causes enormous crop loss throughout the major tea-growing countries of Asia including India, Sri Lanka, Indonesia, China, and Japan causing a yield loss of 40% globally [10]. The incidence and severity of blister blight depend on the nature of the tea cultivar, geographical, and environmental conditions of the tea growing areas. The percent yield loss of made tea due to blister blight incidence across the major tea producing countries is represented in Figure 3b. Some of the most susceptible tea cultivar prone to blister blight infections are UPASI-1 and UPASI-3 (Assam), UPASI-9 and UPASI-15 (China), and UPASI-17 and TRI-2025 (Cambod), BSS-1, etc. [11]. Here is a comprehensive discussion on the incidence of blister blight disease in different countries, the causal organism, disease cycle, epidemiology, severity, and different approaches employed for the control of blister blight.
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1.1 History
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Balidon in his book ‘Tea in Assam’ has indicated the prevalence of blister blight disease on wild indigenous tea in Assam shortly after the beginning of tea cultivation during 1863 [12]. Shortly afterwards Peal in 1868 recognized the existence of blister blight disease of tea and Sir George Watt was the first to report the disease symptoms in Assam in the year 1895. [9, 13]. Later the confirmation of causative pathogen of blister blight as Exobasidium vexans was reported by Massee, the Mycologist of Kew Botanical garden in 1898 based on samples sent by Dr. Watt from Upper Assam, India [14]. In 1908, a sudden outbreak of the disease occurred in Darjeeling, West Bengal, India, and was then subsequently reported from Formosa (Taiwan) in 1912, Japan in 1922, and Taiwan in 1938 [15, 16, 17]. This was followed by the emergence of the disease in southern India in August 1946, wherein blister blight incidence was first reported in tea estates of Mundakayam and Peermade valley in Kerala. The disease soon spread in the West-Northwest and Southwards direction to Anamallais, other tea estates of Kerala, Nilgiris, Wynaad, and Chikmagalur in Karnataka, due to the effect of South West and North East monsoon winds, thereby affecting the entire tea growing regions of southern India [18]. Later on, the disease was reported from Sri Lanka in 1947 [19], from Sumatra and Java in Indonesia in 1949 [20], Nepal in 1948, East Pakistan in 1951, Thailand in 1953, and South Vietnam and Cambodia in 1959 (CMI, 1970). Hence, blister blight has eventually become a devastating tea disease throughout all the major tea plantations of Asian countries including India, Sri Lanka, Bangladesh, Cambodia, China, Indonesia, Japan, Malaysia, Nepal, Taiwan, Thailand, and Vietnam.
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1.2 Blister blight disease symptoms, pathogen, and life cycle
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The causal organism of blister blight disease Exobasidium vexans is known to be an obligate biotrophic fungus with no alternate host completing its entire life cycle in tea (Table 2). The pathogen mainly attacks young, succulent, and tender harvestable leaf and shoot thereby inflicting an enormous effect on the quality and quantity of consumable tea production. The pathogen reproduces through basidiospores which are commonly known to get dispersed by wind. The basidiospores germinated upon lodging on the surface of susceptible tea leaf surfaces under a humid atmosphere with a minimum relative humidity of 80%. The infection is facilitated by the formation of infection peg from appressoria either directly penetrating the cuticle of host tissue or penetration through stomata [21]. The first apparent sign of infection appears in young leaves in the form of pink translucent spots which are considered as the first stage of blister blight infection and are visible after three days of fungal penetration. The spots enlarge along with the leaves and approximately reach a diameter of 3–12.5 mm. In the second stage, the enlarged spots develop into white and velvety convex blister lesions on the lower surface of tea leaf, and on the upper surface, the area with blister lesions becomes sunken resulting in concave depression [22]. The disease progress to its third stage characterized by the curling of the infected tea leaf, browning of blister lesions, and consequently necrosis of the infected leaf tissue. The life cycle of blister blight disease is represented in Figure 4. During the off-season, the pathogen is reported to survive on these necrotic leaf parts which facilitate the infection to occur in the subsequent season under favorable climatic conditions. In a study was carried out to detect the survival of E. vexans during the off-season the presence of basidiospores in the atmosphere was reported indicating the active state of the pathogen throughout the year. However, the spore concentration being very low (10 spores/m3) the basidiospores failed to sporulate [23]. Under favorable climatic conditions, the pathogen completes its life cycle within 11 days, although it is reported at times extend to 28 days depending on the prevailing climatic conditions. Owing to the short span of the life cycle, multiple generations of the pathogen are completed within a single cropping season. For the development of blister blight infection sporulation to germination takes place in 4 h to 5 days, germination to penetration takes 4–9 days, penetration to the appearance of visible symptoms takes 3–10 days, development of mature blister and subsequent sporulation takes 11–28 days [11].
The taxonomic position of E. vexans as described by Massee [14].
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Figure 3.
(a) Climatic condition influencing blister blight disease, (b) blister blight mediated crop loss (%) across major Asian countries [NER: North east region].
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Figure 4.
Disease cycle of blister blight along with life cycle of causal organism Exobasidium vexans.
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A histological study of blister blight disease on tea leaves provided insights into the cellular alteration of host tissue during infection. The study revealed that during the first stage of infection, the enlargement of the translucent spots is a result of hypertrophy as the size of the cells in the mesophyll layer of the infected leaf was substantially higher as compared to the healthy leaf. In the lower epidermis of infected tea leaves with mature blisters, the development of hymenium was prominent in the second stage. This disrupts the lower epidermis completely and gets filled with networks of intercellular hyphae which subsequently develop into basidia that bear basidiospores. However, in tea leaves with blister infections localized in veins, the proliferation of hymenium was apparent in both the lower and upper epidermis. This results in the disruption of the sclerenchyma layer in the vein thereby rupturing xylem and phloem resulting in leaf curling and necrosis of infected leaf in the third stage. The hymenium consisting of bundles of hyphae on maturity forms the clavate to cylindrical basidia (46.98–86.42 μm × 4–5 μm) with normally two and rarely three to four sterigmata [24]. The basidiospores of E. vexans are formed at the apex of these sterigmata and two nuclei from the basidium pass into spore via fission. The basidiospores are hyaline and elliptical and measure 7–15.5 X 2.3–4.5 μm when observed under a microscope. It has been reported that 10,000 basidiospores are produced per mm2 of the blister lesion while the mature blister lesion can produce up to two million basidiospores in 24 hrs [25, 26]. Although the basidiospores are single-celled when immature, three to four septa are reported to form during germination [27]. In a recent study, stages of basidiospore germination were reported under in vitro conditions (Figure 5). The basidiospores were found to germinate on agar surface 4 h post-incubation followed by germ tube growth from either one or both ends. The spores were initially observed to be aseptate and at a later stage, as many as four transverse septa were formed. At 8 h post-incubation, the formation of hyphae was observed that differentiated into branches to form a complex network of hyphae [28]. So far, the identification of E. vexans is being carried out by studying the basidiospore morphology as discussed above, and blister blight disease is identified symptomatically. Molecular based identification remains a major challenge as very few sequences for the ITS region of E. vexans have been deposited in NCBI to date. This urges the development of a specific molecular barcode for the identification of E. vexans. E. vexans being an obligate biotrophic fungus, establishing in vitro culture to study the pathogen is another major challenge. In this context, Sundstrom reported that thiamine is a significant supplement in culture media [29]. This was followed by the use of media based on natural substrates for the growth of E. vexans under laboratory conditions [30, 31]. Also, vitamin B5 and calcium pantothenate has been indicated as necessary supplements to maintain in vitro culture of E. vexans for a period of up to 48 h [32]. Viable in vitro culture of E. vexans was achieved up to four-five weeks from an optimization study on three different basal media. The carbon source was found as the most significant parameter for czapek dox and v8 juice while tea leaf extract for potato dextrose along with optimal temperature and pH range to be 25–27°C and 7–8 respectively [28].
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Figure 5.
Representative image of dorsal (a) and ventral (b) surfaces of blister blight infected tea leaf (Ananda tea estate, Lakhimpur, India). Different phases of germination of E. vexans basidiospores on agar: Basidiospores (c); germ tube growth from single end of basidiospores (d); germ tube growth from both ends of basidiospores (e); and hyphal growth of E. vexans with branching (f). Infected leaf part with extensive hyphal growth (g–h) and formation of appressorium (i) (scale: 10 μm, gt-germ tube, br-branching of hyphae, hy-hyphae, app-appresorium) [adapted with permission from Chaliha et al. 2020].
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1.3 Epidemiology, occurrence, and disease severity
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The weather condition plays a very important role in the epidemiology and severity of blister blight disease. Low temperature, high humidity, cloudy condition with moderate rainfall has been found to play a profound impact on the development of pathogen and disease incidence. As such the incidence of the disease is most favored in monsoon season and facilitated with relative humidity (RH) of more than 80% and availability of water on the leaf surface. In a study carried out by Huysmans (1952), blister blight incidence was recorded with a 5-day average of RH of greater than 83%. On the other hand, Homburg (1953) studied that RH below 80% over 5 days was unfavorable to blister infection. Venkata Ram has reported that the optimum period of leaf wetness to facilitate infection was 11 h and the maximum infection occurred at 13 h [33]. The requirement of moisture content for the germination of basidiospore is reported to be provided by approximately 0.1-inch rain per day while the optimal growth temperature was recorded to be 20–25°C with a maximum tolerance limit of 34°C [29, 34]. Sporulation was found to be inhibited at a temperature greater than 35°C and a temperature of 32°C was reported to be lethal for the basidiospores of E. vexans [35]. The incidence of blister blight disease is inversely related to the period of sunshine. Visser et al. in 1961 found a reduction of blister blight disease with an average of 3.5 h of sunshine per day for 5 days at a stretch [34]. Following this report, an exercise of cutting shade trees to medium height for allowing penetration of sunlight to the tea canopy was practiced in Sri Lanka [36]. In a different study, the UV-B (290–320 nm) component of sunlight was found to reduce the sporulation in blister thereby decreasing the number of spores at the end of the disease cycle [37]. Variation in the nature of spores and sporulation behavior was also observed in basidiospores developed under adverse climatic conditions. The basidiospores produced during unfavorable months were found to be thick-walled that failed to germinate. Also, the atmospheric spore count was less during these months. However, in tea plantations close to the ravine that recorded low temperature and high humidity, blister blight disease was noticed during the unfavorable months. Also, tea plants in these areas experienced surface wetness of 15 h which is ideal for blister blight disease incidence [23]. The optimal climatic condition influencing blister blight disease (Figure 3a) and percentage crop loss in major Asian countries (Figure 3b) is depicted in Figure 3.
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The spore liberation in the air over blister blight infested tea plantations follows a diurnal rhythm and resembles a nocturnal pattern of spore discharge of other basidiomycete pathogens. The maximum liberation of basidiospores was found to occur between midnight and 4.00 am [38]. The spore deposition on tea plants was found to be directly proportional to the number of spores in the atmosphere. However, the difference in spore deposition in different bushes was observed with higher spore deposition in susceptible hosts [39].
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Blister blight disease, being a foliar disease, directly affect the quality and quantity of consumed tea. Severe disease incidence has been recorded after pruning of tea plantations owing to the abundance of young and tender leaf and stem. Also, during infection of tender stem the entire shoot withers and falls along with the curled infected leaf making it unusable for plucking [33]. As such along with enormous yield loss a quality deterioration below 35% disease threshold level is imposed due to blister blight infection [10, 40]. The percentage of crop loss varies with the geographical condition of different countries. In Sri Lanka, Loos reported 50% crop loss in tea plantation without protection, and 33% in plantations protected with copper fungicide [41]. Indonesia reported a loss of ~10 million kg of tea which is 20–25% between 1951 and 1952 [16]. In southern India, during the initial years of blister blight infection, enormous crop losses were observed with an annual loss of about 18 million kg of tea between 1948 and 1952 [42]. North-east India reported, crop loss up to 24% due to blister blight infection, the infection occurring mostly in the hilly region. Darjeeling has been reported with the worst effected tea plantations with blister blight, owing to the favorable climatic conditions. The onset of the disease has been recorded in June with the starting of monsoon and reaches its severity in August till October. In Assam, India blister blight incidence was associated with early rain in February and reaches its severity in the month of March–April [43].
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Blister blight infection results in significant degradation of quality in made tea owing to changes in biochemical characteristics [44, 45]. Gulati in 1999 carried out the analysis of biochemical parameters in diseased leaf. In the infected leaves catechin content, total phenols, nitrogen, chlorophyll, amino acids, and polyphenol oxidase activity was recorded in decreasing concentration in comparison to healthy leaf. In orthodox tea processed from infected tea leaves theaflavins, caffeine, catechin polymer thearubgins, and aroma components were significantly found in reduced concentration [46]. Tea shoots with blister infection were also reported with a decrease in catechin content, flavor component 2-phenyl ethanol, and enzyme activity of prephenate dehydrase [47].
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1.4 Blister blight disease control
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Considering the severity of blister blight disease and its related agronomic and economic losses control of the disease is of utmost necessity. Various control measures have been adopted for the protection of tea plantations against the disease of which the use of therapeutic approaches at the field scale started around 40 years ago [48]. Over the last few years, the various control measures adopted against blister blight disease can be categorized into cultural, chemical, biological, and host tolerance approaches. Different studies have been carried out concerning control measures against E. vexans infection. An overall representation of various approaches in controlling blister blight disease is shown in Table 3. Details of these approaches and their application at field scale are discussed below.
Disease inhibition (%) with various control measures.
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1.4.1 Cultural practice
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\nE. vexans mainly infect young succulent harvestable leaves which are normally plucked for commercial tea production. The cultural practice of hard plucking also known as fish leaf plucking and early pruning has been employed in tea plantations to reduce the severity of blister blight infection. Eden (1947) reported that hard plucking practice after every two to three months in a year causes no major damage to the tea bushes. However, the long-term hard plucking can result in decreasing crop yield as the bushes get weakened and become susceptible to the attack of mite [57]. Also, tea plantations after hard plucking get delayed and irregular with the growth of new foliage, and as such the tea plantations look worn out [41]. Severely infected tea plants are pruned immediately to control blister blight diseases. Pruning was carried out during hot dry weather to ensure the growth of foliage in the period when the disease presented no danger. However, this resulted in the sunscald damage of the stems that developed into cankers [58]. In line with cultural practices, pruning of shade trees is also rehearsed to allow sunlight to fall on tea bushes as long-term sunlight is reported to inhibit the germination of basidiospores of E. vexans [59]. UV-B solar radiation component has been reported to play a significant role in the natural regulation of blister blight disease. The study has shown that removal of UV-B component from sunlight falling on tea bushes resulted in an increasing number of blisters formation while on the other hand, complete sunlight reduced the number of sporulating blisters post 62 h of inoculation. This proves that prolong durations of sunlight can have a negative impact on completing several generations within the same cropping season [37].
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1.4.2 Chemical control
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The importance of chemical control of blister blight disease and the use of economically feasible chemical therapeutics dates back to 1960 when the disease incidence was recorded in southern India. Protectant fungicides, eradicant fungicides, and systemic fungicides are used as foliar sprays against blister blight disease. Bordeaux mixture and copper oxychloride are the two most commonly used protective fungicide formulations. The acceptance level of the use of copper in tea leaves to control blister blight was set at 150 ppm (Lamb, 1950) as copper-based fungicides also possess collateral damage of phytotoxicity and release of copper residues to environment causes human health hazard, effect soil microflora, and marine population. The formulation of copper oxychloride was able to control blister blight disease at a usage rate of 0.21 Kg metallic copper per hectare. The concentration of copper at 50% wettable powder was used in copper oxychloride formulations [59, 60, 61]. Eradicant fungicide nickel chloride hexahydrate was found effective in controlling blister blight disease by antisporulant activity. The reduction in infection was achieved from 84–24% post 3 weeks of treatment and up to 13% after 5 weeks. However, the treatment with nickel chloride was found severely phytotoxic which rejected its use as a potent fungicide [50]. Owing to the phytotoxicity and collateral health hazard from chemical fungicides, organic fungicides were introduced in Sri Lanka, Indonesia, and southern India. However, the disease resistance efficacy was lower in comparison to copper-based fungicides. Also, the high cost related to the processing of organic fungicides discarded its use for blister blight control [25, 60, 62, 63]. Two common brand names for organic fungicides used for blister blight control are Daconil and Difolatan [64].
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Conventionally, around 26 rounds of spraying of these fungicides are carried out at 7-days intervals during the disease season to control blister blight incidence. However, since climatic conditions play a significant influence on the severity of blister blight incidence the spraying interval of fungicides differs from region to region. In Indonesia and Sri Lanka an extended period of spraying based on sunshine hours at a specific period of the disease season mediated the control of disease at the economic threshold. On the other hand, control of blister blight disease was not achieved even after a 7-day spraying interval in southern India [65]. Systemic fungicides are often used against plant pathogens owing to its sustained control of plant for example, blister blight could be controlled in southern India by administering pyracarbolid (Sicarol) over 3 weeks. This treatment exhibited strong antisporulant activity reducing the sporulation in mature blisters, while eradicating the latent blister lesions. Also, plant growth was found to be stimulated with the use of pyacarbolid [33]. In a different study with systemic fungicides ergosterol biosynthesis inhibiting (EBI) fungicides tridemorph, bitertanol, hexaconazole, and propiconazole were studied for its effect on physiological parameters of the tea plant and controlling blister blight disease in southern India. EBIs were found with antisporulant activity with a significant reduction in spore size, viability, and inhibited spore germination except for tridemorph treatment. As such, inhibition in spore germination reduced the viability of spore which mediated the reduction in spore load thereby controlling blister blight incidence. The effectiveness of treatment lasted for 7 days, with a reduction of the occurrence of the disease by half relative to untreated plants. Additionally, the EBIs were found with a positive effect on the physiological parameters of the tea plant. The stomatal conductance, total chlorophyll, carotenoids, and photosynthetic rates were found to be induced with EBIs treatment along with an increase in biometric parameters like dry weight and shoot length [49]. In North-East India, 2–3 rounds of systemic fungicides like propiconazole or hexaconazole has been used at 5% EC @ 1:1000 as a foliar spray at 14 days interval to control blister blight infection (Tea board of India).
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1.4.3 Biological control
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Besides showing appreciable control of blister blight disease with chemical therapeutics, the related phytotoxicity and health hazard have initiated the approach of biological control of blister blight disease. The use of biological control agents like Trichoderma harzianum, Serratia marcescens, Gliocladium virens, Bacillus subtilis, and Pseudomonas fluorescens have been studied against blister blight disease [66, 67, 68, 69]. However, the use of these bioformulations was not found efficient in controlling blister blight disease. Plant growth-promoting rhizobacteria Pseudomonas and Bacillus were tested for the control of blister blight disease. Foliar application of Pseudomonas fluorescens Pf1 bioformulation at 0.5% concentration weekly showed appreciable efficiency against blister blight disease and the lowest mean disease index of 16.06% was achieved which was at par with the chemical fungicide (14.57%). Reduction in disease incidence was achieved for two seasons and the treated tea plants were found with an induced accumulation of defense enzymes peroxidase, polyphenol oxidase, b-1,3-glucanase, chitinase, phenylalanine ammonia-lyase, and phenolics as compared to control. As such, PGPR mediated induced systemic resistance of tea plants against blister blight infestation [48]. Tea phylloplane bacteria have been isolated and screened for their inhibitory action against blister blight disease, of which isolate identified as Ochrobactrum anthropi and designated as BMO-111 was found efficient for the biocontrol of E. vexans. Foliar application with BMO-111 at 15 days intervals up to 120 days recorded a reduction in disease incidence. Treatment with BMO-111 resulted in 73.4% protection, compared to 64.7% protection with chemical treatments. An inhibitory effect of basidiospore germination and antifungal effectivity was achieved against E. vexans. However, the mechanism of action is still unknown and urges further investigation [52]. Although the majority of studies in this context have been carried out in India, a recent study in Indonesia reported the potential use of soil bacteria (Azoto II-1) and three endophytic (Acinetobacter sp., Endo-5, Endo-65, and Endo-76) bacteria against blister blight incidence. The bacterial suspension was to the soil of infected tea plantations at a dose of 2 l ha−1, applied six times at a 1 week interval. The bacterial formulations were observed to control blister blight intensity only up to 2 weeks after treatment. However, the reduction of disease was not significant as compared to control. Also, the treatment was not accompanied by an increase in yield of fresh shoots except for Acinetobacter sp. that showed a 17.26% increase in fresh shoot yield [53]. Thus, in the context of biological control of blister blight disease, bioformulation of microbes and its commercialization at the field scale urges further studies for the screening of beneficial microbes that may be applied for efficient management of blister blight disease.
\n
\n
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1.4.4 Host resistance against blister blight disease
\n
In the context of blister blight disease, a variety of tea cultivars have been reported that are found to be resistant to E. vexans infection. Various studies have been carried out on the morphological characters of these cultivars and the various blister blight defense-related enzymes and pathways have been analyzed to study the nature and basis of resistance. A study was carried out with blister blight resistant clone SA-6 and a susceptible clone TES-34. Both the clones were studied for anatomical differences such as cuticle and epidermal thickness, stomatal length and breadth, palisade tissue, and quantification of epicuticular wax. They have reported that the resistant clone SA-6 showed a higher amount of epicuticular wax, the high thickness of cuticle with greater stomatal frequency when compared to TES-34. Upon pathogen infection, PR proteins are reported to induce systemic acquired resistance in plants. As such, the study also analyzed the difference in the production of PR protein chitinase and have found higher constitutive expression of chitinase in resistant clone [70]. In a recent study with these cultivars, SA6, and TES34, the difference in gene expression against blister blight infection was accessed for chitinase, glucanase, phenylalanine ammonia-lyase, and genes in the flavonoid pathway. The relative intensity of the expression of these genes was found to be higher in the resistant cultivar SA6 in comparison to susceptible cultivar TES34. Also, the expression of these pathogenesis-related genes was found to increase along with each successive stage of blister blight infection [71]. A similar study was conducted recently on the biochemical characterization of resistant tea cultivar AV-2 and susceptible cultivar B-157. Secondary metabolites phenol and proanthocyanidin content were reportedly higher in the resistant clone AV-2. As such, the inferred resistance may be attributed to the antifungal properties of phenol and free radical scavenging activity with the chelation of transition metals by proanthocyanidins. Also, a higher concentration of hydrolyzing enzyme acid phosphatase, peroxidase, catechol oxidase, and superoxide dismutase occur in AV-2 cultivar as compared to B-157 [72].
\n
In line with understanding the basis for resistance of tea cultivar against blister blight disease transcriptome profiling of two cultivars P-1258 (resistant) and T-78 (susceptible) have been carried out to identify defense-related transcripts particularly in a resistant cultivar. cDNA-AFLP mediated the screening of differentially expressed candidate transcripts which mainly showed homology with an acyl-CoA binding protein, zinc finger family protein, ubiquitin, and proline-rich protein that were upregulated after infection. Suppression subtractive hybridization-based transcriptome analysis resulted in a comprehensive study of transcripts induced in resistant cultivar P-1258 after infection. The induced contigs showed similarity to proteins such as ubiquitin family protein, an iron–sulfur cluster scaffold protein, short-chain dehydrogenase, ribulose-1,5-bisphosphate carboxylase/oxygenase small subunit, thioredoxin, pathogenesis-related proteins (chitinase, endo-glucanase, beta-glucosidase, wound-induced protein, protease inhibitor, thaumatin-like protein, cystatin, blight associated protein p12, aspartic proteinase) and proteins with a function in defense signal transduction pathway (serine/threonine-protein kinase, oxo-phytodienoic acid reductase, mitogen-activated protein kinase, leucine-rich repeat transmembrane protein kinase, salicylic acid-binding protein, calcium ion binding or calmodulin-related protein, hydrogen peroxide-induced protein, chitin-inducible gibberellin responsive protein, calreticulin). qRT-PCR based expression analysis of the genes showed greater than two-fold upregulation in P-1258 when compared to T-78 post-infection. Hence, the expression profiling mediated the molecular characterization of resistant tea cultivar involved in developing possible systemic acquired resistance against E. vexans infection [73]. To provide further insights into the molecular mechanism of host resistance against blister blight disease, a similar study of transcriptome profiling was carried out with tea cultivars SA6 (resistant) and Kangra-Asha (susceptible) at different stages of blister blight disease. In SA6 cultivar at stage 1 (post 24 h infection), salicylic acid metabolism and secondary metabolite biosynthetic processes were recorded to be enhanced. At stage 2 (7 dpi) hydrogen peroxide metabolic processes, cellular metabolism, and ion transport metabolism-related genes were found to be enriched in resistant variety. As such, hydrogen peroxidase activity suggests efficient scavenging of free radicals enabling restricted penetration/germination of spores inside the host tissue thereby conferring resistance. At stage 3 of infection (14 dpi) increased expression of phenylpropanoid (PAL) and aromatic compound category of gene synthesis probably lead to the synthesis of antimicrobial compounds that might have protected intercellular hymenium development in SA6 cultivar. It was interesting to note that at stage 4 (20 dpi) during necrosis of blister infected part Jasmonic acid-mediated signaling pathway genes are enriched in resistant variety SA6 along with induced monooxygenase and ACC oxidase activity and ethylene production [10]. Here, the induction of both SA and JA signaling pathways again urges to examine the hemibiotrophic existence of the pathogen which is otherwise reported to be biotrophic. In a recent study presence of hypothetical proteins (HPs) was identified and assigned with novel putative defense-related functions in a resistant cultivar of tea SA6 against blister blight disease. The HPs proteins were functionally categorized into LRR, WRKY, NAC, chitinases, and peroxidases. Additionally, different pathways playing a significant role in SA6 resistance against blister blight were annotated based on the KEGG database including plant-pathogen interaction, biosynthesis of secondary metabolites, metabolic pathways, amino sugar, and nucleotide sugar metabolism, and phenylpropanoid biosynthesis which have probably [74].
\n
Plants use various defense mechanisms to shield themselves from infection by pathogens. The cell wall itself acts as an insulation against invading pathogens. Pathogens invading plants breach the cell wall by releasing enzymes and the products get accumulated in the apoplastic region. These are termed elicitors and are capable of activating a complex array of defense signaling called pathogen triggered immunity. These elicitors also mediate the induction of systemic acquired resistance (SAR) in plants. In the last few years, various biotic and abiotic elicitors have been tested and found to mediate SAR in plants against the pathogen. Two chemical elicitors acibenzolar-S-methyl and salicylic acid were tested for their efficiency to induce SAR against blister blight in tea. Plants treated with 0.1% ASM provided 40.8% protection against blister blight. Salicylic acid was used at 250 ppm to achieve significant induction of resistance. Tea plants treated with elicitors were recorded with an induced level of β-1,3-glucanase, phenylalanine ammonia-lyase, and peroxidase activity thereby conferring resistance against blister blight disease [54]. In a similar study treatment of tea plant with abiotic elicitor calcium chloride (CaCl2), found to induce activities of defense enzymes like phenylalanine ammonia lyase (PAL), polyphenol oxidase, peroxidase, and b-1,3-glucanase along with a higher accumulation of total phenolics, thaumatin, cinnamate 4-hydroxylase, flavonoid 30-hydroxylase when compared to control plants [55]. In this context, the use of chitosan as elicitors in tea plants to provide resistance against blister blight disease has been tested and the possible mechanism of resistance has been analyzed. Chitosan solution applied as a foliar spray at 0.01% concentration and 15 days interval reduced blister blight incidence for two seasons. The induced resistance was found to be facilitated by nitric oxide (NO) signaling and the level of total polyphenol content and expression of defense-related enzymes (peroxidase, polyphenol oxidase, phenylalanine ammonia-lyase, and b-1,3 glucanase) was induced [56].
\n
Genetic improvement of tea has been made possible with transgenic technology started from the year 2000. Gene technology and the development in agrobacterium-mediated transformation mediated the incorporation of a foreign gene into crop plants for the development of cultivar with resistance against various diseases. Agrobacterium-mediated transformation was used to develop a disease-resistant variety of tea against blister blight with the introduction of Solanum tuberosum class I chitinase gene into tea genome. Plant selectable marker hygromycin phosphotransferase (hpt) gene was used to confer hygromycin resistance. 12 tea plantlets were confirmed with stable integration of transgene that showed resistance against blister blight disease tested in a detached leaf infection assay [75]. In a study carried out by the same group transgenic tea cloned with Solanum tuberosum endo-1,3-beta-D-glucanase using Agrobacterium-mediated transformation technique showed significant resistance against blister blight disease. Upregulation of pathogenesis-related (PR) genes like PR3 (chitinase I) gene and PR5 (thaumatin-like protein) gene was recorded in the transgenic tea plantlets which can be attributed to the resistance against blister blight disease [76]. A similar study was carried out for co-transformation of tea genome with LBA4404 pCAMBIA 1301-Chi (carrying potato class I chitinase gene expression cassette) as reported in Singh et al. (2015) and LBA4404 pBI121-Def (carrying mung bean defensin gene expression cassette) together. Although resistance against blister blight was achieved with the co-transformation the resistance was better with Solanum tuberosum class I chitinase gene introduced transgene.
\n
\n
\n
\n
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2. Conclusion
\n
Widespread research has been carried out so far on understanding the incidence of blister blight infection and on its control measures for the survival of the tea industry. However, to date identification of the pathogen is being carried out symptomatically and morphologically. In the context of molecular identification of E. vexans a few sequences for the ITS region are found in the NCBI database which urges to carry out a more detailed study on the development of molecular barcode-based identification. Also, very little is known so far about the genome and transcriptome profile of the pathogen which could be a basis for understanding the molecular mechanism behind the pathogenesis of E. vexans. Understanding the molecular basis of pathogenesis of E. vexans would likely mediate control measures to be applied more specifically and efficiently. In line with control measures against blister blight, most of the studies reported so far have worked on defense responses and resistance mediated by tea cultivars and hence urges further studies to elucidate the molecular pathogenesis cycle of the pathogen. As such, from the pathogen point of view, there is a significant gap in understanding blister blight disease and a thorough analysis of the pathogen is likely to address the various associated challenges.
\n
\n
Acknowledgments
\n
Author CC would like to acknowledge DST, Govt. of India for her DST INSPIRE Junior Research Fellowship (IF-150964).
\n
Conflict of interest
The authors declare that they have no known potential conflict of interest.
\n',keywords:"tea, blister blight, Exobasidium vexans, Basidiospore, defense, control",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/74600.pdf",chapterXML:"https://mts.intechopen.com/source/xml/74600.xml",downloadPdfUrl:"/chapter/pdf-download/74600",previewPdfUrl:"/chapter/pdf-preview/74600",totalDownloads:141,totalViews:0,totalCrossrefCites:0,dateSubmitted:"September 8th 2020",dateReviewed:"December 4th 2020",datePrePublished:"December 28th 2020",datePublished:null,dateFinished:"December 28th 2020",readingETA:"0",abstract:"Tea is one of the most popular beverages consumed across the world and is also considered a major cash crop in countries with a moderately hot and humid climate. Tea is produced from the leaves of woody, perennial, and monoculture crop tea plants. The tea leaves being the source of production the foliar diseases which may be caused by a variety of bacteria, fungi, and other pests have serious impacts on production. The blister blight disease is one such serious foliar tea disease caused by the obligate biotrophic fungus Exobasidium vexans. E. vexans, belonging to the phylum basidiomycete primarily infects the young succulent harvestable tea leaves and results in ~40% yield crop loss. It reportedly alters the critical biochemical characteristics of tea such as catechin, flavonoid, phenol, as well as the aroma in severely affected plants. The disease is managed, so far, by administering high doses of copper-based chemical fungicides. Although alternate approaches such as the use of biocontrol agents, biotic and abiotic elicitors for inducing systemic acquired resistance, and transgenic resistant varieties have been tested, they are far from being adopted worldwide. As the research on blister blight disease is chiefly focussed towards the evaluation of defense responses in tea plants, during infection very little is yet known about the pathogenesis and the factors contributing to the disease. The purpose of this chapter is to explore blister blight disease and to highlight the current challenges involved in understanding the pathogen and pathogenic mechanism that could significantly contribute to better disease management.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/74600",risUrl:"/chapter/ris/74600",signatures:"Chayanika Chaliha and Eeshan Kalita",book:{id:"10113",title:"Diagnostics of Plant Diseases",subtitle:null,fullTitle:"Diagnostics of Plant Diseases",slug:null,publishedDate:null,bookSignature:"Dr. Dmitry Kurouski",coverURL:"https://cdn.intechopen.com/books/images_new/10113.jpg",licenceType:"CC BY 3.0",editedByType:null,isbn:"978-1-83962-516-9",printIsbn:"978-1-83962-515-2",pdfIsbn:"978-1-83962-517-6",editors:[{id:"264297",title:"Dr.",name:"Dmitry",middleName:null,surname:"Kurouski",slug:"dmitry-kurouski",fullName:"Dmitry Kurouski"}],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 History",level:"2"},{id:"sec_2_2",title:"1.2 Blister blight disease symptoms, pathogen, and life cycle",level:"2"},{id:"sec_3_2",title:"1.3 Epidemiology, occurrence, and disease severity",level:"2"},{id:"sec_4_2",title:"1.4 Blister blight disease control",level:"2"},{id:"sec_4_3",title:"1.4.1 Cultural practice",level:"3"},{id:"sec_5_3",title:"1.4.2 Chemical control",level:"3"},{id:"sec_6_3",title:"1.4.3 Biological control",level:"3"},{id:"sec_7_3",title:"1.4.4 Host resistance against blister blight disease",level:"3"},{id:"sec_10",title:"2. Conclusion",level:"1"},{id:"sec_11",title:"Acknowledgments",level:"1"},{id:"sec_14",title:"Conflict of interest",level:"1"}],chapterReferences:[{id:"B1",body:'\nMandal, A. B., Basu, A. K., Roy, B., Sheeja, T. E., & Roy, T. (2004). Genetic Management for Increased Tolerance to Aluminium and Iron Toxicities in Rice—A Review.\n'},{id:"B2",body:'\nHarbowy ME, Balentine DA, Davies AP, Cai Y. 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Effect of aqueous vitamin B on the growth of blister blight pathogen, Exobasidium vexans. Songklanakarin Journal of Science & Technology. 2012;34(6):601-606\n'},{id:"B33",body:'\nVenkata Ram CS. Calixin, a systemic fungicide effective against blister blight (Exobasidium vexans) on tea plants. Pesticides. 1974;\n8\n:21-25\n'},{id:"B34",body:'\nVisser T, Shanmuganathan N, Sabanayagam JV. The influence of sunshine and rain on tea blister blight, Exobasidium vexans Massee, in Ceylon. Annals of Applied Biology. 1961;\n49\n(2):306-315\n'},{id:"B35",body:'\nVenkata Ram CS, Chandra Mouli B. Systemic fungicides for integrated blister blight control. UPASI Tea Sci. Pep. Bull. 1976;\n33\n:70-87\n'},{id:"B36",body:'\nArulpragasam PV, Addaickan S, Kulatunga SM. Recent developments in the chemical control of blister blight leaf disease of tea - effectiveness of EBI fungicides. S.L.J. Tea Sci. 1987;\n56\n:22-34\n'},{id:"B37",body:'\nGunasekera TS, Paul ND, Ayres PG. 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Journal of Applied Microbiology. 2013;\n114\n(1):209-218\n'},{id:"B53",body:'\nFauziah F, Setiawati MR, Pranoto E, Susilowati DN, Rachmiati Y. Effect of indigenous microbes on growth and blister blight disease of tea plant. Journal of Plant Protection Research. 2019:529-534\n'},{id:"B54",body:'\nAjay D, Baby UI. Induction of systemic resistance to Exobasidium vexans in tea through SAR elicitors. Phytoparasitica. 2010;\n38\n(1):53-60\n'},{id:"B55",body:'\nChandra S, Chakraborty N, Chakraborty A, Rai R, Bera B, Acharya K. Abiotic elicitor-mediated improvement of innate immunity in Camellia sinensis. Journal of Plant Growth Regulation. 2014;\n33\n(4):849-859\n'},{id:"B56",body:'\nChandra S, Chakraborty N, Panda K, Acharya K. Chitosan-induced immunity in Camellia sinensis (L.) O. Kuntze against blister blight disease is mediated by nitric-oxide. Plant Physiology and Biochemistry. 2017;\n115\n:298-307\n'},{id:"B57",body:'\nDe Weille GA. Blister blight Exobasidium vexans in tea and its relationship with environmental conditions. Netheriand J. Agile. Sti. 1960;\n8\n(3):183-210\n'},{id:"B58",body:'\nVenkata Ram, C.S. (1974b). Pruning for rejuvenation. Planters1 Chron. 69, 279-282.\n'},{id:"B59",body:'\nVisser T, Shanmuganathan N, Sabanayagam TV. Blister blight control in 1957 with respect to fungicidal formulation application rates and yield. Tea Quart. 1958;29:9-20\n'},{id:"B60",body:'\nJayaraman V, Venkataramani KS. Control of blister blight in tea in southern India. The 1956 field trials. Planters\' Chron. 1957;52:35-39\n'},{id:"B61",body:'\nJayaraman V, Venkata Ram CS. Control of blister blight of tea in southernIndia. The 1958 field trials. Ann. Rep. UPASI Sci. Pep. Tea Sect, for. 1959;1958-59:28-35\n'},{id:"B62",body:'\nLaoh JP. Fungiclden proeven bij. blister blight (Exobasidium vexans) op. thee. Arch, Voor. Thee Cult. 1955;\n19\n:1-9\n'},{id:"B63",body:'\nVenkata Ram CS. Application of nickel chloride to tea plants (Cameltla &inw6i6) and control of blister blight. Current Science. 1960;30:57-58\n'},{id:"B64",body:'\nVenkata Ram CS. Systemic control of Exobasidium vexans on tea with 1, 4-Oxathiin derivatives. Phytopathology. 1969;\n59\n:125-126\n'},{id:"B65",body:'\nRam CV, Mouli BC. Interaction of dosage, spray interval and fungicide action in blister blight disease control in tea. Crop Protection. 1983;\n2\n(1):27-36\n'},{id:"B66",body:'\nPremkumar R. Report of the plant pathology division. Annual Report of UPASI Tea Research Foundation pp. 2001:32-33\n'},{id:"B67",body:'\nPremkumar R. Report of the plant pathology division. Annual Report of UPASI Tea Research Foundation pp. 2002:35-36\n'},{id:"B68",body:'\nPremkumar R. Report of the plant pathology division. Annual Report of UPASI Tea Research Foundation pp. 2003:38-39\n'},{id:"B69",body:'\nBalasubramanian S, Parathiraj S, Haridas P. Effect of vermicompost based Trichoderma (Vermiderma) on the recovery of pruned bushes and on the control of certain diseases in tea (Camellia sinensis (L) O. Kuntze). J. Plantation Crops. 2006;\n34\n(3):524-528\n'},{id:"B70",body:'\nJeyaramraja PR, Pius PK, Manian S, Meenakshi SN. Certain factors associated with blister blight resistance in Camellia sinensis (L.) O. Kuntze. Physiological and Molecular Plant Pathology. 2005;\n67\n(6):291-295\n'},{id:"B71",body:'\nNisha SN, Prabu G, Mandal AKA. Biochemical and molecular studies on the resistance mechanisms in tea [Camellia sinensis (L.) O. Kuntze] against blister blight disease. Physiology and Molecular Biology of Plants. 2018;\n24\n(5):867-880\n'},{id:"B72",body:'\nHazra A, Dasgupta N, Sengupta C, Kumar R, Das S. On some biochemical physiognomies of two common Darjeeling tea cultivars in relation to blister blight disease. Archives of Phytopathology and Plant Protection. 2018;\n51\n(17-18):915-926\n'},{id:"B73",body:'\nBhorali P, Gohain B, Gupta S, Bharalee R, Bandyopadhyay T, Das SK, et al. Molecular analysis and expression profiling of blister blight defense-related genes in tea. Indian Journal of Genetics and Plant Breeding. 2012;\n72\n(2):226\n'},{id:"B74",body:'\nSingh G, Singh G, Seth R, Parmar R, Singh P, Singh V, et al. Functional annotation and characterization of hypothetical protein involved in blister blight tolerance in tea (Camellia sinensis (L) O. Kuntze). Journal of Plant Biochemistry and Biotechnology. 2019;\n28\n(4):447-459\n'},{id:"B75",body:'\nSingh HR, Deka M, Das S. Enhanced resistance to blister blight in transgenic tea (Camellia sinensis [L.] O. Kuntze) by overexpression of class I chitinase gene from potato (Solanum tuberosum). Functional & Integrative Genomics. 2015;\n15\n(4):461-480\n'},{id:"B76",body:'\nSingh HR, Hazarika P, Agarwala N, Bhattacharyya N, Bhagawati P, Gohain B, et al. Transgenic tea over-expressing solanum tuberosum endo-1, 3-beta-D-glucanase gene conferred resistance against blister blight disease. Plant Molecular Biology Reporter. 2018;\n36\n(1):107-122\n'}],footnotes:[],contributors:[{corresp:null,contributorFullName:"Chayanika Chaliha",address:null,affiliation:'
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