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Section 1

Tea, Tea Drinking and Varieties

Chapter 1. The Tea Plants

Chapter 2. Green Tea

Chapter 3. White Tea

Chapter 4. Black Tea

Chapter 5. Pu-erh Tea

Chapter 6. Tea Flavanols

Chapter 7. Analysis of Antioxidant Compounds in Different Types of Tea

Chapter 8. Cultivar Type and Antioxidant Potency of Tea Product

Chapter 9. Objective Evaluation of the Taste Intensity of Tea by Taste Sensors

Chapter 10. Green Tea (Cv. Benifuuki) Powder and Catechins Availability

Chapter 1

The Tea Plants

Botanical Aspects

F.N. Wachira¹,³, S. Kamunya¹, S. Karori², R. Chalo¹ and T. Maritim¹

¹Tea Research Foundation of Kenya, Kericho, Kenya

²Department of Biochemistry, Egerton University, Egerton, Kenya

³ASARECA, P.O. Box 765, Entebbe, Uganda


AFLP    amplified fragment length polymophism

EC    (−)-epicatechin

ECG    (−)-epicatechin gallate

EGC    (−)-epigallocatechin

EGCG    (−)-epigallocatechin gallate

EST    expressed sequence tag

F1    first filial generation

GA    gallic acid

GABA    gamma aminobutyric acid

GC    (−)-gallocatechins

GCG    (−)-gallocatechin gallate

IPGRI    International Plant Genetic Resources Institute

LSI    late acting prezygotic gametophytic self incompatibility

PPO    polyphenol oxidase

RAPD    random amplified polymorphic DNA

RFLP    restriction fragment length polymorphism

SSR    simple sequence repeat

STS    sequence tag site

TF    theaflavins

TI    Terpene Index

TR    thearubigins


The cultivated plant species Camellia sinensis ((L.) O. Kuntze) is the source of the raw material from which the popular tea beverage is processed. The species is now cultivated commercially in Asia, Africa and South America. Major producers of the crop include China, India, Kenya, Sri Lanka and Indonesia (Table 1.1). Kenya is currently the largest single exporter of tea (Table 1.2). Although the crop is cultivated in many countries, there are several different types of tea plant, each with its own identifiable character and potential for unique cup quality. Because of this diversity, it is important that the different types of tea plant can be told apart and be classified. Classification, in the biological sense, is the ordering of plants into a hierarchy of classes. The product is an arrangement or system of classification designed to express inter-relationships and to serve as a filing system. The term ‘classification’, however, is often used for both the process of classifying and for the system which it produces.

TABLE 1.1 World Production of Tea (Metric Tons) and Percent Share

TABLE 1.2 World Exports of Tea (Metric Tons) and Percent Share

Classification in Camellia

The tea plant (Camellia sinensis) from which the beverage tea is processed, is placed in the genus Camellia. The genus has over 200 species and is largely indigenous to the highlands of Tibet, north eastern India and southern China (Sealy, 1958). Sealy (1958) classified the genus into 12 subgeneric sections, one of which (Thea) contains species of cultivated tea. However, in his monograph Sealy recognized a group of 24 inadequately known species which he called ‘Dubiae’ (Dubious). In their work, Chang and Bartholomew (1984) not only translated the 1981 monograph of the genus Camellia by H.T. Chang but also included publication of new taxa and moved many species treated by Sealy to different sections. They divided the genus into four subgenera (sub groups), i.e. Protocamellia, Camellia, Thea and Metacamellia, and twenty sections (Figure 1.1).

FIGURE 1.1 Summarized Schematic Diagram Showing Species Relationships within Genus Camellia .

Taxonomy of the genus Camellia has been complicated by the free hybridization between species, which has led to the formation of many species hybrids (Chuangxing, 1988). Similarly, most species are unavailable to scientists for study. Genetic relationships and taxonomy has therefore remained controversial and recent interest has seen the discovery of many new species and a revision of taxonomic relationships (Chuangxing, 1988; Lu and Yang, 1987; Tien-Lu, 1992). Tea is, however, the most important of all Camellia spp. both commercially and taxonomically. Though the other non-tea Camellia’s are not widely used to produce the brew that goes into the cup that cheers, several species, e.g. C. taliensis, C. grandibractiata, C. kwangsiensis, C. gymnogyna, C. crassicolumna, C. tachangensis, C. ptilophylia, are used as sources of tea-like beverages in parts of China, which indicates that the economic potential for beverage production from additional underutilized species is very great (Tien-Lu 1992; Chang and Bartholomew, 1984). Seed oil from several species including C. fraterna, C. japonica and even C. sinensis are important sources of cooking oil in China. In addition, many Camellia species are of great ornamental value.

At the species level, tea taxonomy failed to attract much attention and interest once the species of economic importance were identified. It continues to be a low-priority area in most tea research programs. The array of hybrids available which might suggest unrestricted introgression of many species of Camellia and tea compound the taxonomic jigsaw. Several minor taxa have been treated as conspecific with major taxa, although more recently accumulated evidence has shown that these minor taxa have no natural distribution and are derived from hybridization events involving different species (Parks et al., 1967; Uemoto et al., 1980). The taxonomic affinities of most interspecific and intraspecific hybrids are unknown, but could provide clues to the evolutionary organization of the tea gene pool. Information on taxonomic characteristics, genetic diversity and biogeography of Camellia in living collections are scantily documented, though vital in identifying sources of desirable genes (Banerjee, 1992).

Tea was initially classified as Thea sinensis by Linnaeus (Linnaeus, 1753). Following the discovery of its economic importance, and the subsequent extensive collection of indigenous teas from the forests contiguous to the upper Assam–Burma–Tibet borders, two distinct taxa were identified and classified by Masters (1844) as Thea sinensis, (the small-leaved China plant) and Thea assamica (the large-leaved Assam plant). For a long time, Thea and Camellia were considered as separate genera (Fujita et al., 1973) and some authors even considered Camellia to be a ‘section’ under the genus Thea (Roberts et al., 1958; Barua and Wight, 1958).

Another group of authors (Sealy, 1958; Barua, 1965) considered that CameIlia and Thea were so much alike in morphological, anatomical and biochemical features that the classification schemes proposed above were unrealistic. According to them, the apparent difference in leaf pose, patina and pigmentation was a part of the total variation in leaf features. Wight (1962) considered Thea to be synonymous with Camellia and the name Camellia prevailed. Thus, today tea is botanically referred to as Camellia sinensis (L.) O. Kuntze, irrespective of species-specific differences. Camellia sinensis is classified under section Thea along with 18 other species (Figure 1.1).

At the species level, several intergrades resulting from unrestricted intercrossing between disparate parents have been documented, but have not been assigned the status of separate species (Sealy, 1958). However, three distinct tea varieties have been identified on the basis of leaf features like size, pose and growth habit. These are the China variety, Camellia sinensis, var. sinensis (L.); the Assam variety, Camellia sinensis var. assamica (Masters) Kitamura; and the southern form also known as the Cambod race, C. assamica ssp. Lasiocalyx (Panchon ex Watt). The three main taxa can be differentiaed by foliar, floral and growth features (Tables 1.3 and 1.4) and by biochemical affinities (Sanderson, 1964; Robert et al., 1958; Hazarika and Mahanta, 1984; Ozawa et al., 1969; Fujita; et al., 1973; Owuor et al., 1987). It is common to find the three different varieties (China, Assam and Cambod) referred to as separate species, namely, Camellia sinensis, C. assamica and C. assamica ssp. Lasiocalyx, respectively (Bezbaruah, 1976). Research has shown that cultivated tea is an out-crosser with an active late-acting pre-zygotic gametophytic self incompatibility (LSI) system (Wachira and Kamunya, 2005a; Muoki et al., 2007). Because of its out-breeding nature and, therefore, high heterogeneity, most cultivated teas exhibit a cline extending from extreme China-like plants to those of Assam origin. Intergrades and putative hybrids between C. assamica and C. sinensis can themselves be arranged in a cline of specificity (Wight, 1962). Indeed because of the extreme hybridizations between the three tea taxa, it is debatable whether archetype (original) C. sinensis, C. assamica or C. assamica ssp. lasiocalyx still exist (Visser, 1969). However, the numerous tea hybrids currently available are still referred to as Assam, Cambod or China depending on their morphological proximity to the main taxa (Banerjee, 1992).

TABLE 1.3 Criteria Used for Differentiating Two Major Tea Varieties and Sub-Varieties of Camellia sinensis

TABLE 1.4 Types of Tea Differentiated on the Basis of Foliar Characteristics

1 = Extreme China

2 = Typical between Assam and China

3 = Typical between Assam and China

4 = Extreme Assam

5 = Close to 2

(Scheme as used by Hadfield (1974))

Recent revisions of taxonomy within section Thea have led to identification and description of additional varieties of C. sinensis, namely var. waldenae (Hu) (Chang and Bartholomew 1984), var. dehungensis (Chuangxing, 1988) Ming, and var. publimba (Chang) (Tien-Lu, 1992). In Taiwan, a new subspecies of wild tea C. sinensis ssp. buisanensis (Sasaki) Lu and Yang, has been described (Lu and Yang, 1987).

Wood and Barua (1958) and Cannell et al. (1977) have raised doubts regarding whether or not the existing tea populations have resulted from natural hybridizations between the three main taxa only, or have also involved other Camellia species. Ackerman (1973) described interspecific crosses between tea Camellias and C. kissi and C. sasanqua. The presence of some tea clones with brick red or purple pigmented leaves may be an indication of possible species hybridization (personal observation; Kerio et al., 2012). According to Wight and Barua (1957), the presence of punctate leaf forms in some tea hybrids provides further evidence of the natural hybridization of both C. sinensis and C. assamica with C. irrawadiensis (a non-tea Camellia) which has similar punctations. The hybrids described, however, produce commercially low-quality tea. Based on this experience, introgression from wild species into the gene pool of tea may not always be beneficial, as this could adversely affect quality attributes (Banerjee, 1992). Indeed, though most non-tea Camellias, e.g. C. irrawadiensis, C. taliensis, C. reticulata, C. pitardii, C. saluensis, C. sasanqua, C. japonica, C. lutescens, C. kissi, C. caudata, C. rosaeflora, C. hongkongensis, C. cuspidata, superficially resemble tea, they either lack caffeine or produce weak infusions which may not pass as quality tea. Directed species hybridization may, however, break this rule with the development of high-quality interspecific hybrids. Indeed presently, many interspecific hybrids of tea and other Camellia species have been developed, for example with C. japonica (Bezbaruah and Gogoi, 1972; Takeda et al., 1987). C. taliensis (Fuchinoe, 1975), C. irrawadiensis (Bezbaruah, 1975), C. sasanqua (Fuchinoe, 1975), C. kissi (Bezbaruah and Saikai, 1977) and C. caudata (Bezbaruah and Saikai, 1977). Special focus has, however, been made on two taxa, C. irrawadiensis and C. taliensis. These two taxa closely resemble C. sinensis variety assamica in leaf and growth characteristics but lack caffeine, with C. irrawadiensis containing an analog of caffeine, theobromine. Because of the aforementioned, the liquors of C. irrawadiensis and C. taliensis lack the quality of tea. All F1 interspecific hybrids created between these two species and C. sinensis have also failed to produce a tea of commercially acceptable quality. Most of these have very low caffeine levels though when one of the C. sinensis X C. irrawadiensis, hybrids was back-crossed to C. sinensis, it gave high-quality, high-yielding progeny (Bezbaruah, 1987). The prospect of using interspecific hybrids for improving stress tolerance traits, e.g. cold hardiness, drought resistance, disease and pest resistance, and specific characters in biochemical components, without necessarily impairing quality, therefore exist and need to be exploited further. Nevertheless, it is generally accepted that only three taxa, C. assaica, C. sinensis, C. assamica ssp. lasiocalyx, have contributed to the gene pool of cultivated tea. Thus, the term ‘tea’ is often used to include the above taxa and their hybrids. The contribution of the recently described varieties, waldenae, dehungensis, publimba and subspecies buisanensis, to cultivated tea is as yet unknown.

Methods of Distinguishing Types of Tea

One of the key elements in classification is description. Description is the orderly technical recording of characteristics of a group, particularly the morphological ones. A diagnosis is a short comparative description, stating only those characters which differentiate the taxon from its closest allies in the same rank. Characterization calls for development of suitable descriptors for elucidating differences between tea plants. Ideally such descriptors should not be affected by plant development and growth environment.

Morphological Traits

Though the tea plant can grow to become an under tree or even a tree, cultivated plants are maintained as a low bush in a continuous vegetative phase of growth by cyclic pruning every three to five years to form a plucking table. Because of this cultural practice, it has become essential that vegetative characteristics be used to describe and differentiate tea taxa. However, because of its outbreeding nature and high heterogeneity, most of its vegetative, biochemical and physiological characteristics show continuous variation and high phenotypic plasticity (Purseglove, 1968; Wickremasinghe, 1979; Wickremaratne, 1981; Banerjee, 1988; Magoma et al., 2000; 2003). Despite this, leaf macromorphological features, e.g. leaf color, size and pose; leaf angle and leaf area index; have been widely used as descriptors in tea taxonomy, although chemotaxonomy (Takeo, 1983; Owuor et al., 1987; Magoma et al., 2000; Kerio et al., 2012) and molecular (DNA) taxonomy (Matsumoto et al., 1994; Wachira et al., 1995, 1997, 1999; Paul et al., 1997; Kaundun and Matsumoto, 2002), have also been applied.

The International Plant Genetic Resources Institute (IPGRI) has published a standardized descriptor list for tea (IPGRI, 1997). This list provides an international format for characterization in a universally understandable ‘language’ for tea genetic resources data. Traditionally, leaf and floral morphology, and growth habit have been the more important criteria for assigning taxonomic categories within Camellia. The standardized descriptor list of the IPGRI also includes such novel methods as chemical profiling, e.g. for catechin content and Terpene Index; biochemical markers, e.g. isoenzymes; cytological markers, e.g. chromosome number, meiosis chromosome associations, identified and sequenced genes, etc.; and the more esoteric molecular marker fingerprint profiles, e.g. restriction fragment length polymorphisms (RFLP), random amplified polymorphic DNA (RAPD), amplified fragment length polymorphism (AFLP), simple sequence repeats (SSRs), sequence tag sites (STS), and expressed sequence tags (EST), etc.

The vegetative characteristics generally used in assigning taxonomic categories in tea include: leaf size (leaf length, leaf breadth), ratio of leaf length to breadth, internode length, length and girth of bud, petiole length, ratio of apical length, angle between leaf tip and axis, leaf margin, shoot density, etc. Although these related characteristics may overlap and show continuous variation, restricting their usefulness in characterization somewhat, they continue to be widely used by tea growers and scientists the world over. Based on growth habit and leaf features, the major varieties of tea have been distinguished as the China and Assam varieties, as shown in Table 1.3. This classification scheme identifies two separate sub-varieties of the China variety: f. parviflora and f. macrophylla. It is difficult to distinguish between f. parviflora and f. macrophylla, except that the former possesses extremely small (1.5–6.0 cm long; 1.2–2.0 cm wide) and the latter relatively large (4.0–14.0 cm long; 2.0–2.5 cm wide) leaves (Bezbaruah, 1976). It is doubtful if these two taxa still exist in existing commercial tea populations, and whether or not they can be set apart from C. sinensis var. sinensis. They possibly represent a part of the total variation in leaf features in the chinary type of tea (Bezbaruah, 1976). Purseglove (1968) mentions a third variety, C. sinensis var. macrophylla Makino from Japan. It is a large-leaved triploid (2x = 3n = 45), and gives a bitter decoction.

Based on foliar characteristics, Hadfield (1974) recognizes five types of tea, each with distinct and quantifiable foliar characteristics. Though these types are not separate taxa, the variations in their foliar attributes can be related to either extreme China or extreme Assam plants (Table 1.4). Hadfield (1974), quoting Purseglove (1968), did not consider the involvement of any variety other than Assam and China in accounting for the variation in leaf characteristics in the five types of tea.

The grouping of tea into the erect, small-leaved China variety and the horizontal, broad-leaved Assam variety was rather subjective because plants with intermediate leaf characteristics could not always be assigned to either of these two varieties. Only on the basis of leaf angle, for example, can tea be classed as erectophile (leaf angle <50°, planophile (leaf angle >70°, or oligophile (leaf angle 50–70°). While China and Assam types generally cover the erectophile and planophile plants respectively, the oligophiles are far too distinct to come under any of the two main varieties. Despite this limitation, leaf features continue to be widely used to classify tea and to study the extent of variation between cultivated taxa.

An example is presented in which some vegetative traits are evaluated in a collection of Kenyan tea clones (Table 1.5). In the example, the Assam and Cambod varieties had larger leaves and leaf pose angle than the China variety. The tetraploid clones in particular had very large leaves. The non tea Camellias, i.e. C. irrawadiensis and C. japonica, had leaves of similar dimensions to those of the Assam variety. Using hierarchical cluster analysis and the leaf size traits (length and breadth) of the first four leaves (Nos. 1–4), it was possible to distinguish the China variety clones from the rest (Table 1.6). The TRFK clone Dwarf, which has extreme China characteristics, clustered alone. In this assay, the C. irrawadiensis and C. japonica clustered with the Assam varieties. A similar analysis using the leaf pose angles and leaf internodes gave different clusters with the former, distinguishing the Cambod clone TRFK 301/3 and the latter again distinguishing the TRFK clone Dwarf. A combination of all the traits did not improve the resolution of clustering; this meant that leaf size alone was fairly useful in discriminating varieties of Kenyan tea. This has also been confirmed by the use of other statistical analysis methods, such as principal component analysis (Wachira, unpublished). This further demonstrates the power of leaf traits in differentiating tea taxa.

TABLE 1.5 Vegetative Traits in Some Kenyan Tea Clones

Superscript, leaf base angle.

+ploidy, no of chromosomes, i.e. diploid = 2x = 2n = 30, triploid = 45, tetraploid = 60

∗Camellia irrawadiensis

∗∗Camellia japonica

TABLE 1.6 Cluster Membership of Tea Clones Using Average Linkage (Between Groups) Analysis

+All the combined traits

∗C. irrawadiensis

∗∗C. japonica

Leaf anatomical differences and particularly the variation in distribution and morphology of sclereids in leaf lamina have also been useful for differentiating tea varieties. Cambod clones have numerous sclereids compared to Assam and China clones. Similarly, sclereids in Cambod tea have wider lumen when compared to those of the other varieties (Barua and Wight, 1958). Leaf trichome hairs have also been used to identify some tea taxa (Amma, 1986). Stomata, leaf anatomical features and stomatal conductance have also been used to characterize tea clones (Ng’etich and Wachira, 2003).

Unlike foliar features, floral morphology can also provide reliable diagnostic criteria for differentiating various tea taxa and classifying them into discrete groups. However, because tea in production is maintained at a vegetative phase of growth by cyclic pruning, these features are not widely used and are not considered in this article.

Cytological Markers

The main varieties of tea are diploids (2n = 30), and their chromosome structures are also comparable; even C. irrawadiensis conforms to 2n = 30. The chromosome number of several ‘wild’ tea (e.g. C. caudata and C. kissi), is also of 2n = 30 as in cultivated tea. This constancy in diploid chromosome number might suggest a monophyletic origin of all tea species, including perhaps C. irrawadiensis and other Camellia species. Deviations from normal chromosome number are however more common in Japanese Camellia taxa than in taxa from other areas. C. sasanqua is a hexaploid or tetraploid (2n = 90 or 2n = 60), C. sinensis var. macrophylla is a triploid (2n = 45) and C. rosaeflora, a tetraploid with 2n = 60.

Cultivated tea forms a stable polyploidy series. Natural triploids, tetraploids and aneuploids have been sampled in tea populations in Japan, India and Kenya, but are reportedly present in very low number (Wachira and Kiplangat, 1991; Wachira, 1994). Generally, rooting ability, leaf size and dry-weight of polyploids as well as total polyphenol content are higher than those of diploids (Wachira, 1994: Wachira and Ng’etich, 1999; Magoma et al., 2000). Pollen viability and fertility of triploids are, however, usually poor; with tetraploids being more fertile than the triploids, but less than the diploids (Wachira and Kiplangat, 1991). Studies carried out on different tea cultivars have revealed that chromosome complements of tea comprise of near metacentric to submetacentric chromosomes (Bezbaruah, 1975; Wachira et al., 1999). Nucleolar number in tea has been demonstrated to correspond to multiples of the somatic cell number and is a good marker for ploidy in the species (Wachira and Muoki, 1997). Cytological investigations in tea have, however, been restricted mostly to determination of chromosome number rather than explaining the cytological basis of species differentiation. For example, it is not known whether, in view of similarities in chromosome morphology and identical chromosome numbers, species differentiation in tea started from the same basic genome, or different species and taxa had diverse origins. Detailed studies on chromosome morphology in different hybrids might be helpful in elucidating the cytological relationship.

Chemical/Biochemical Traits

Chemical profiling has also been used to characterize species within the genus Camellia (Nagata, 1986). Tea has been demonstrated to contain characteristic compounds, such as caffeine, catechins and theanine (Table 1.7). Only species of section Thea in the entire genus of Camellia contain galloyled catechins such as (−)-epicatechin gallate (ECG) and (−)-epigallocatechin gallate (EGCG) and the non protein amino acid, theanine. Anthocyanin-rich tea has also been described in Japan and Kenya (Terahara et al., 2001; Kerio et al., 2012). Eugenol glycosides, sasanquin and fluorescent flavonoid sulfates have been used to study introgression of C. japonica traits into C. sasanqua (Parks et al., 1981).

TABLE 1.7 Leaf Biochemicals in Plants from the Genus Camellia

Caf, caffeine; Thb, theobromine; Thea, theanine; EC, (−)epicatechin; (+)C, (+)-catechin; EGC, (−)-epigallocatechin; ECG, (−)-epicatechin gallate; EGCG, (−)-epigallocatechin gallate; SQN, sasanquin; –, <0.01%; I, 0.01–0.3%; II, 0.3–1.0%; III, >1.0%; N.D., not detected, D., detected. (After Nagata, 1986.)

The IPGR descriptor list recognizes some chemical parameters in black processed tea as highly discriminative. These include theaflavin and thearubigin contents and fractions, tea quality type and the terpene index (TI). Leaf terpeniods have successfully been used to characterize closely related Camellia species (Takeo, 1983; Nagata, 1986; Owuor et al., 1987) though at higher levels of classification, they are less useful owing to convergence and reticulate evolution. Owuor et al. (1987) used the terpene index (TI) to distinguish between some Kenyan tea clones. Though some overlap was noted, it was established that the Assam types had TI close to 1.0 while the China types had TI near zero. Although Hazarika and Mahanta (1984) characterized Assam, China and Cambod tea based on variations in chlorophylls a and b and four carotenoids (neoxanthine, violaxanthine, B-carotene and lutein), these characteristics were unstable and varied with season, making them ineffective in large-scale systematic studies of tea. Magoma et al. (2000) used the ratio of dihydroxylated to trihydroxylated catechins to delineate different varieties of tea. Studies of this nature are important to establish the affinities of hybrids to the major taxonomic categories.

Total tannin content and caffeine have also been used to discriminate tea clones (Takeda, 1994). In a study comparing tea germplasm from India, Bangladesh, Sri Lanka, Taiwan, China and Japan, Takeda (1994) found that the Japanese and Chinese germplasm were low in their total tannin content and caffeine content. Studies in Kenya have also revealed the Japanese tea as low in total polyphenols when compared to Kenyan germplasm (Wachira and Kamunya, 2005b). In Japan, low-tannin-content cultivars are selected for production of green tea. In Kenya, tannin and caffeine profiles have been widely used to characterize the Kenyan tea germplasm. Wide variations in tannin and caffeine content have been observed in the Kenyan tea germplasm (Table 1.8). Further, tea cultivars can also be classified based on their capacity to undergo auto-oxidation as a consequence of different polyphenol oxidase (PPO) activity. In Kenya, a non-fermenting (PPO-deficient) tea plant has been selected (personal observation).

TABLE 1.8 Summarized Data of Average Tannins Content in a Collection of Kenyan Tea Clones

Molecular Markers

The emergence of molecular techniques has provided a rapid and efficient means of examining accumulated genetic differences in the tea gene pool without interference of the environment. Molecular markers have provided an important set of descriptors which are able to differentiate between genotypes, variety types and even different species of the genus Camellia. These markers are highly discriminative and can even distinguish genotypes that cannot be distinguished using morphological traits. They are useful for the determination of differentiation and relatedness within cultivated tea. In a study that profiled three NADP-linked dehydrogenase isozymes, Magoma et al. (2003) revealed that tea clones could be accurately partitioned according to their phylogenetic origins. The development of DNA markers has further improved the ability to identify tea clones in commercial use and more so those used as breeding stocks (Wachira et al., 1995; Paul et al., 1997; Wachira et al., 2001; Singh and Ahuja, 2006). They also provide a measure for accumulated genetic variability (Wachira et al., 2001). Molecular markers have also been demonstrated to be taxonomically informative at the species and genus level. Phylogenetic relationships in Camellia which have been constructed using molecular data have been similar to those obtained using morphological and biochemical affinities (Wachira et al., 1997). Molecular markers, however, provide a greater degree of resolution. Moreover, molecular tools have been utilized to generate genetic linkage maps of tea and to identify genes responsible for important agronomic and biochemical traits (Matsumoto et al., 1994; Hackett et al., 2000; Kato et al., 2000; Kaundun and Matsumoto, 2002; Chen et al., 2005a, b; Ma and Chen et al., 2006; Kamunya et al., 2010).


Cross Compatibility

Compatibility between species within the genus Camellia has been used to construct species relationships (Takeda, 1990). Several Camellia species hybridize with tea (Table 1.9). The relationships constructed from species affinities are however not useful for description of individual tea varieties and clones.

TABLE 1.9 Summary of Results of Crosses between Tea and Some Allied Species in the Genus Camellia

With tea (C. sinensis) as female species. (Summary after Takeda, 1990.)

Summary Points

• Tea is classified in the subgeneric group Thea of the genus Camellia.

• Several members of the genus Camellia freely interbreed among themselves.

• The numerous interspecific hybrids between members of Camellia have complicated taxonomy within the genus.

• The tea beverage is largely processed from Camellia sinensis ((L.) O. Kuntze).

• In China, several other species of Camellia are used to make tea-like beverages.

• Cultivated tea plants are largely differentiated by leaf morphophysiological traits.

• The three major cultivated forms of tea include the China variety (Camellia sinensis var. sinensis (L.)); the Assam variety (Camellia sinensis var. assamica (Masters) Kitamura) and the Cambod race (C. assamica ssp. Lasiocalyx (Panchon ex Watt)).

• The small-leafed China variety is predominantly cultivated in China and Japan for the production of unaerated green tea, while the large-leafed Assam variety is cultivated in India, Sri Lanka, Africa and Argentina for the production of aerated black tea.

• The three predominant varieties of tea differ in their morphological, physiological and leaf biochemical traits as well as in their molecular profiles.

• New emerging consumer requirements may necessitate tea breeding for high or low functional components, e.g. in catechins, flavanoids, theanine, caffeine, theobromine, Gamma Amino Butyric Acid (GABA), b-carotene, astringency, aroma, etc.


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Chapter 2

Green Tea

The Plants, Processing, Manufacturing and Production

Selena Ahmed¹ and John Richard Stepp²

¹Department of Biology, Tufts University, Medford, MA, USA

²Department of Anthropology, University of Florida, Gainesville, FL, USA


Green tea is sourced from Camellia sinensis (L.) O. Kuntze (Theaceae), the same species from which white, oolong, black and pu-erh teas are derived. The various tea types are classified on the basis of their processing, and the associated oxidation and fermentation levels which influence taste and aroma profiles. Green tea is a minimally oxidized and non-fermented tea. Catechin polyphenols are the primary compounds responsible for the claimed health benefits of green tea, including its antioxidant and anti-inflammatory properties. Caffeine contributes to green tea’s stimulant properties, while the amino acid theanine contributes to its relaxing properties. Significant variation of catechin and caffeine content occurs between green tea types depending on environmental growth conditions and processing. Most often, green teas are sourced from the sinensis variety of the tea plant while black teas are sourced from the assamica variety. China is the center of origin of green tea production. Today, green tea is produced in over 20 countries in tropical, sub-tropical and temperate regions. It is the most widely consumed beverage after water, due to its health, sensory, stimulant, relaxing and cultural properties. The quality of green tea is impacted by numerous factors involved in cultivation, harvest, processing, storage and preparation that influence chemistry, taste, aroma, morphology and bioactivity of tea leaves.

The Plants: Botanical Classification and Distribution

Botanical Identification

The tea plant was first described taxonomically in 1753 by Carl Linnaeus in Species Plantarum. He referred to the tea plant as Thea and later refined the species into black tea (Thea bohea) and green tea (Thea viridis). By the early 1900s, taxonomists recognized that green and black tea were both from the same species, Camellia sinensis (L.) O. Kuntze.

Camellia sinensis is an evergreen tree or shrub. It has yellow-white flowers and long, serrated leaves. Flowers are axillary, solitary or up to three in a cluster. They are 2.5–3.5 cm in diameter and have six to eight petals. The outer petals are sepaloid and the inner petals are obovate to broadly obovate. There are numerous stamens 0.8–1.3 cm in length. Young leaves have short white hairs on their underside and young branches are grayish yellow and glabrous. Current year branchlets are purplish red. Terminal buds are silvery gray and sericeous. Petioles are 4–7 mm in length, pubescent and glabrescent. Leaf blades are elliptic, oblong-elliptic or oblong. Seeds are brown, subglobose and 1–1.4 cm in diameter. Flowering of Camellia sinensis occurs from October through February and fruiting occurs from August to October (Min and Bartholomew, 2007).

The tea plant belongs to the Theaceae botanical family. The Theaceae (Mirb. ex Ker Gawl.) is part of the order Ericales (Bercht. and J.Presl) in the Asterid branch of eudicots (Angiosperm Phylogeny Group III, 2009 (Haston, et al., 2009)). This family includes approximately 600 species (Tsou, 1998). The only autapomorphy (a distinctive anatomical feature unique to a terminal group) that identifies Theaceae is a specialized pseudopollen (Tsou, 1998).


There are four varieties of Camellia sinensis including Camellia sinensis var. assamica, Camellia sinensis var. sinensis, Camellia sinensis var. dehungensis, and Camellia sinensis var. pubilimba. Commercial tea is primarily produced from the former two varieties, Camellia sinensis var. assamica, or the broad-leaf variety of the tea plant, and Camellia sinensis var. sinensis, or the small-leaf variety of the tea plant. Green tea is most often sourced from the small-leaf variety because it has a sweeter taste than the broad-leaf variety that is usually used for black and pu-erh tea production. However, both varieties can be used to produce white, green, yellow, red, black and pu-erh teas.

The Yunnan-Guizhou Plateau is the center of differentiation of the two main commercial Camellia sinensis varieties. Tea plants occur as the broad-leaf variety south of 25° N in the Yunnan-Guizhou Plateau and as the small-leaf variety north of this latitude, with various hybrid types being found in between (Ming, 1992). Tea plants of the broad-leaf variety are quickly growing plants of either tree or bush form, suitable for warm, tropical and sub-tropical environments (Banerjee, 1992). Individuals of the small-leaf variety grow as shrubs or dwarf plant forms and are more suitable for colder, temperate climates. Differentiation between the two commercial Camellia sinensis varieties is further defined by matK and rbcL nucleotide sequence polymorphisms of the chloroplast locus (Katoh et al., 2003; Stoeckle et al., 2011).


The tea plant naturally grows in the understory of broad-leaved evergreen forests at altitudes between 100–2,200 m. The native tea growing area, or ‘tea belt’, encompasses southwestern China (Yunnan, Sichuan, Guangxi and Guizhou provinces), northern Laos, northern Vietnam, Myanmar, Cambodia and northeastern India. Some forest-growing tea plants are also found outside of the tea belt in eastern China, Japan, southern Korea, Thailand and Taiwan. The center of diversity of Camellia sinensis is in the Upper Mekong River region of Yunnan Province of southwestern China (Chen and Pei, 2003; Chen et al., 2005). Today, tea plants are primarily found in terraced landscapes in the tea belt and other tropical, subtropical and temperate regions where tea cultivation has been introduced, including Sri Lanka, Indonesia, central Africa, Turkey, Argentina and Russia. Tea plants are found between latitudes 42° N and 30° S in areas that fit the specific ecophysiological requirements of tea cultivation (see Cultivation section below).

Tea seeds were initially dispersed through waterways such as the Mekong River and by human exchange. It is unlikely that animals disperse tea seeds over long distances because of the seeds’ large size. The broad-leaf Camellia sinensis var. assamica is prevalent across the tea belt and the small-leaf Camellia sinensis var. sinensis prevails in eastern China, Taiwan, Japan, India’s Darjeeling and tea-growing countries outside of Asia (Ming, 1992). The biogeographic distribution of the Camellia sinensis varieties are a function of adaptation, cultivation history and taste preferences. The small-leaf variety primarily used for green and white tea production is generally cultivated in eastern China, Taiwan and Japan. This variety is also grown in India’s Darjeeling region to produce delicate black teas. The broad-leaf variety is generally cultivated in western China, Myanmar and Assam to produce black, green and pu-erh teas. However, both varieties can be used to produce different tea types and both varieties can be found across these regions. Hybridization occurs where the Camellia sinensis varieties overlap, resulting in a highly heterogeneous botanical product (Takeda, 1990).

The distribution and naming of the two commercial Camellia sinensis varieties is linked to British colonial relations with China and India. Tea became popular with Europeans after their forays in eastern China in the 17th century. At that time, China was the global supplier of tea, exporting the small-leaf variety of the tea plant. The British were forced to find other sources of tea to fulfill demand when Chinese tea became inaccessible during the Opium Wars in the mid 19th century. They turned to the cultivation of tea in India, which soon became the primary source of tea for the British Empire. While India did not commercially produce tea before the 1800s, tea plants grew in forests in Assam and were harvested and used by indigenous communities. Tea specimens that were collected from Assam were distinguished from tea plants from eastern China as a separate variety, the two being respectively termed Camellia sinensis var. assamica and Camellia sinensis var. sinensis. It was later recognized that the assamica variety also grows in forests and indigenous tea production systems in southwestern China where its center of diversity has been identified.

The British East India Company led commercial tea cultivation in India by transferring tea germplasm and technology from China (Murray, 1852). Scottish botanist Robert Fortune was sent to smuggle and transport tea from China to India in 1848 (Fortune, 1853). Fortune used Wardian cases, transportable greenhouses invented by Nathaniel Bagshaw Ward, to transport thousands of tea seeds and seedlings from eastern China to India’s Darjeeling region. The British East India Company also brought trained tea farmers and producers from China to transfer knowledge of tea cultivation and processing. Forest tracts were cleared across Darjeeling and Assam for tea plantations, in which both the broad-leaf and small-leaf varieties were cultivated.


Tea has been cultivated for at least 1,500 years. Cultivation of tea is possible in areas that receive over 120–150 cm of rain annually, and have temperatures of 12–30 °C. Optimum growing conditions are 250–300 cm annual rainfall and average temperatures of 18–20 °C (Williges 2004). Cultivation may occur from sea level to 2,200 meters, with some tea cultivars found as high as 3,000 m. Higher altitudes are often associated with higher tea quality. At least five hours of direct or 11 hours of indirect sunlight daily are required for tea cultivation. Soils must be well drained, sandy, thoroughly aired, deep and nutritious with a healthy layer of humus and low pH. Drought, water logging, excessive heat and frost are harmful for the growth of tea plants and may result in a lower quality product in terms of chemistry, taste, aroma and bioactivity. Tea plants are often raised in controlled nursery conditions or other protected conditions for their first two to four years. They are classified as immature at this time and are not harvested. Once tea plants mature, they are transplanted to the field and are ready for harvest.

Propagation and Breeding

Tea plants cross-pollinate and are self-incompatible (Banerjee, 1992). They are propagated sexually by seeds or asexually by vegetative cuttings of clonal propagules. Grafting may be also used for vegetative propagation of tea plants where seedlings may be used as rootstocks. Tea plants cultivated from seeds generally give lower yields, require less fertilization and produce a heterogeneous stand and product compared to plants propagated by vegetative cuttings. Seed propagation also results in tea plants with an extensive taproot that helps protect soils, prevent erosion and enhance fertility. Historically, tea plantations were created through seeds and some producers and consumers prefer the heterogeneous product yielded by this method.

In contrast, many large-scale producers prefer clonal propagation by vegetative cuttings because of the uniform product yielded by this method. Clonal propagules need greater fertilization compared to tea plants grown from seed because of their lack of a taproot. This means that they are often associated with poor soil because the short root limits soil microbial interactions. Vegetative cuttings were found to require 45–60 kg nitrogen (N), 4–7 kg phosphorus (P), 20–30 kg potassium (K) and 4 kg calcium (Ca) per hectare to yield 4 tons of leaf (Rehm, 1989). In addition, clonal propagules generally require greater agro-chemical inputs and have narrower genetic diversity compared to tea plants grown from seed. Some environmental conditions favor clonal proagules, while others result in greater vulnerability for clonal propagules compared to seed plants (Williges, 2004). Grafting has been used to select for various traits from two different cultivars, and has a benefit over clonal propagation in that the resulting plants have a taproot system similar to plants grown from seed. Taproots help protect soils and maintain fertility. This propagation method has been adapted as an alternative method to enhance fertility, along with other management practices such as intercropping, alley cropping and organic agriculture.

Tea plants are bred for desired traits through selection and hybridization. These focus on morphology, yield, resistance to abiotic and biotic stress, therapeutic properties, taste and aroma. Selection and hybridization have resulted in over a thousand cultivars and landraces with diverse characteristics. For example, for a grafted plant, a rootstock may be selected from a drought-tolerant cultivar and combined with a scion from a high-yielding cultivar with specific chemistry, taste and aroma profiles. The resulting composite plants combine the selected characteristics from both cultivars. Selection has been the main process of breeding tea plants because of the lack of success of controlled hybridization crosses. However, selection is limited by the natural history of the tea plant. Limitations to breeding tea include the perennial nature of the plant with a long gestation period, self-incompatibility, short flowering time of 2–3 months and the long period of time needed for seed maturation – between 12–18 months (Mondal et al., 2004). Selection is further limited by the low availability of mutants with tolerance to specific biotic and abiotic stresses, low success rate of hand pollination, low propagation rate, and differences of flowering time and fruit-bearing capability. More recently, producers have tried to breed tea plants through the development of transgenic plants but this also has had limited success because of inefficient transformation.

Shrub Formation

The majority of tea plants cultivated globally are grown in compact rows and are pruned as roughly rectangular shrubs at a height of 1–1.2 m in what is termed a picking or plucking table (Figure 2.1). Formation of a plucking table serves to expedite harvest and to maximize yields per area. Young tea plants are usually laterally cut approximately 23 cm above the soil to encourage lateral growth and formation of a shrub. Pruning is done throughout the year, between harvest seasons or annually to encourage growth and maintain the form of the plucking table. Deep pruning occurs every four to five years.

FIGURE 2.1 Tea Plucking Tables in Terrace Tea Production System.

Tea plants in terrace plantations are grown in compact rows to optimize yields per area and are maintained as shrubs by pruning at a height of 1–1.2 m for ease of harvest.

Pests and Disease

Tea plants have a number of naturally occurring predators including the short-hole borer beetle (Xyleborus fornicatus), red spider mite (Oligonychus coffeae), coccinellid and staphylinid larvae, lace-wing larvae, and Typhlodromus and Phytoseius mites (Williges, 2004). Some tea pests such as the short-hole borer beetle cause severe damage to the frame of tea plants. For some green tea types, infestation by particular pests is favorable for the sensory properties of the tea products. Insecticides have conventionally been used to control tea pests. Tea plants are also vulnerable to blister blight disease from the fungus Exobasidium vexans. This fungus is eliminated by direct exposure to sunlight and thus shade trees are removed in areas that are susceptible to infestation. Copper-based fungicides are also used to eliminate to blister blight disease.

Tea Harvest: Plucking and Seasons


Most green tea types are produced using young, tender leaves. The leaf unit harvested varies with the specific type of green tea. Many green tea types include the terminal bud, the internodes and one to three leaves immediately below the bud (Figure 2.2). For some other green tea types, the harvested unit consists of a single young leaf. Older leaves are generally not used to produce high-quality green teas because these leaves are rough with an astringent flavor. Tea leaves are plucked by hand or are mechanically harvested by a plucking machine. Many high-quality green teas are hand plucked. Farmers may hand pluck up to 30 kilos of fresh tea leaf per day. Hand plucking is the most labor intensive and expensive process in tea cultivation and has been replaced in many instances by mechanized pluckers to increase labor productivity and decrease labor costs. Leaves are plucked from the same plant at intervals of four days to two weeks.

FIGURE 2.2 Harvested Unit.

The young tender leaves of tea plants are harvested to produce most types of green tea. Many high-quality teas consist of the terminal bud, internodes and two to three leaves immediately below the bud.

Harvest Seasons

The tea harvest calendar is based on the availability of young terminal buds and adjacent leaves of the tea plant and is partly determined by the proportion of daylight and darkness in a day. There are three to four tea harvesting seasons, depending on the latitude and altitude of a tea production system. At latitudes above 16° N and below 16° S, where tea plants receive less than 11 hours of sunlight for over six weeks, tea plants go into dormancy from late fall to the end of winter. Leaf growth resumes in the spring after a period of dormancy. These areas have three harvest seasons including spring, summer or monsoon season, and autumn. The winter dormancy is regarded an important contributor to good quality tea because it encourages the young shoots to release aromatic compounds in the spring harvest season that are not found in teas of later harvests. At latitudes below 16° N and above 16° S, tea plants continue to produce new leaf shoots throughout the year. These areas have four harvest seasons. Harvest seasons may be preceded by pruning tea plants in order to encourage new leaf growth.

In southwest China, tender buds and leaves sprout during three continuous seasons from spring to autumn. Spring starts in early to mid March and runs through mid May; it is regarded as the main harvest season, giving the best-quality tea. The monsoon or rainy season proceeds the spring season and runs through August. Tea leaves are perceived to have the lowest quality during the rainy seasons on the basis of their taste, aroma and health-promoting properties, and farmers may receive half the price for dried leaf during this period compared to the spring season (Ahmed, 2011a). Excessive rainfall leads to soil exhaustion through leaching and possibly impacts the physiological responses of the plant itself. The autumn season, termed the ‘rice flower season’ in parts of southwestern China, follows the rainy season and lasts till early to mid November. Autumn tea is of higher quality than rainy-season tea but lower quality than the spring season. Some farmers report that the spring harvest season has started earlier each year over the past decade because of increased temperatures (Ahmed, 2011a). This may be the result of microclimate variability or global climate change.

Tea plants in India and Japan are also plucked during three harvest seasons. The spring season in India starts in late February to early March and runs through April. Spring tea is characterized as full-bodied, aromatic, slightly bitter and astringent. The pre-monsoon or summer season runs from mid May to mid June or end of June and the monsoon harvest starts early to mid June through August. Monsoon season tea is often not harvested because of its low quality as judged by a mild taste and aroma. The autumn harvest starts in September to the end of October and this tea is characterized as more full-bodied than monsoon tea but less sharp than spring tea. In Japan, the spring season starts in mid April to early May and runs to end of May or early June. Over half the year’s harvest is collected during the spring. The summer harvest starts in mid June to early July and runs to the end of August to early September. Tea leaves are often not harvested during the hottest and most humid parts of the summer season when the quality is the lowest. The fall harvest starts end of September and runs into October.

In Sri Lanka, tea plants are continuously harvested all year round and do not enter dormancy. The topography of the tea garden in Sri Lanka plays a critical role in determining the arrival of the monsoon and the associated tea quality. Tea plants located in cultivation systems in the southwest region are potentially exposed to monsoons from May to August when tea quality is the lowest. Tea quality is highest for winter and spring teas that are produced December to March. Plants located in tea cultivation systems in the northwest region are exposed to monsoons from October to January. The most valued teas from these cultivation systems are produced from May to September before the monsoon season.


A cup of tea prepared from processed leaves tastes, smells and appears notably different than an infusion from fresh tea leaves because of a series of biochemical changes that occur during processing. Fresh tea leaves have a bitter taste and are rich in volatile compounds and aroma precursors. Processing is among the most important factors for flavor and promotes the development of volatile compounds, reduces bitterness, deactivates enzymes, reduces moisture content, and transforms fresh tea leaf into the six different colors used to classify tea: white, green, yellow, blue-green, red and black. The primary steps of green tea processing following harvest include post-harvest spreading out, fixing, rolling, shaping and drying. Upon harvest, tea leaves are promptly processed to transform their flavor and prevent degradation of biochemical profiles and spoilage. On average, four to six kilos of fresh tea leaf are processed into one kilo of dried tea leaf, which contains 1,000–12,000 young shoots.

The cultivation, processing and preparation of green tea as a skilled practice and art developed in China during the 8th century by Buddhist monks, and was disseminated to the public by one monk in particular, Lu Yu, in The Classic of Tea. Buddhist monks were instrumental in spreading the practice of tea production and consumption. They cultivated tea plants in and around their monastery gardens that they would harvest and process. They drank tea during meditation and used it as a tonic to maintain wellbeing. Diversification of tea products expanded during the Ming Dynasty (1368–1644) in China when producers began to experiment with new processing techniques. Today, there are numerous types of green tea, characterized by a range of cultivation and processing methods (Table 2.1).

TABLE 2.1 Examples of Types of Green Tea