Technical Bulletin 4 December, 1965

THE MORPHOLOGY AND VARIETAL CHARACTERISTICS OF THE RICE PLANT

TE-TZU CHANG Geneticist and ELISEO A. BARDENAS Assistant Taxonomist Illustrated by ARNULFO C. DEL ROSARIO Artist-Ilustrator THE INTERNATIONAL RICE RESEARCH INSTITUTE Los Baños, Laguna, The Philippines Mail Address: Manila Hotel, Manila

CONTENTS

Introduction

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Morphology of the Rice Plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Seedling morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Vegetative organs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Culm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Floral organs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Panicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Trade Terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Growth stages of the rice plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glossary of morphologic terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Botanical and Agronomic Traits Useful in Varietal Classification and Identification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seedling characteristics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adult plant characteristics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Classification of cultivated varieties of O . sativa . . . . . . . . . . . . . . . . . . . Mutant Traits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Variations in anthocyanin pigmentation . . . . . . . . . . . . . . . . . . . . . . . . . . Variations in non-anthocyanin pigmentation . . . . . . . . . . . . . . . . . . . . . . Modifications in size and shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Presence or absence of structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Modifications in structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Modifications in chemical composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . Modifications in growth habit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Modifications in other physiological characters . . . . . . . . . . . . . . . . . . . . Glossary of mutant traits and gene symbols . . . . . . . . . . . . . . . . . . . . . . . Acknowledgments ............................................... 12 13 17 17 18 26 27 27 27 27 28 28 28 28 28 29 32 33 36

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Introduction

In describing methods for measuring and recording varietal characteristics, simple and quick operations are preferred to elegant techniques that require precision apparatus. Considerable imWorkers long have recognized the need for uni- portance is given to the economic usefulness of formity in genetic nomenclature of rice. This led the trait under description. Emphasis is also given the International Rice Commission in 1959 to adopt to growth Characteristics of tropical varieties a set of genetic symbols. Comprehensive reviews which constitute the bulk of the rice acreage of of genetic studies and linkage analysis have been the world. It is recognized that many of the propublished. The International Rice Research Insti- posed techniques and methods are inadequate to tute has now assumed the task of monitoring gene describe fully. the enormous variation found in cultivated varieties or to cover the intricate growth symbols. behavior of the rice plant in different environments. Additional studies are needed to improve This publication (1) proposes a set of reason- existing methods. ably definitive terms that adequately describe the Rice workers are urged to adopt the proposed various parts of the rice plant and its processed products, (2) defines varietal characteristics that terms and methods of description, although some are useful in identification and classification, and of them may appear inadequate, and to suggest (3) describes a number of commonly observed ways and means of improvement.

The wide geographical distribution of the rice plant ( Oryza sativa L.) and its long history of cultivation in Asian countries have led to the development of a great diversity of varietal types. Similarly, workers in various rice-growing countries use different terms to designate identical morphological and physiological characters, agronomic traits, gene symbols, and cultural practices. Whereas varietal diversity in germ plasm is desired in rice breeding, variations in nomenclature hinder scientific communication among the workers.

mutant traits in both morphological and genetical terms. In selecting morphological names; terms based on botanical considerations take priority over agronomic terms of extensive usage. Synonyms are also included to provide a basis for concurrent use of terms.

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O. The seminal roots are sparsely branched and persist only for a short time after germination. culms. all of which develop lateral roots. The coleoptile. The length of the axis between the coleoptile and the point of union of the root and culm is called the mesocotyl. blade.4 m. which encloses the young leaves. As other taxa in the tribe Oryzeae. The primary leaf is green and cylindrical and has no blade.. he cultivated rice plant ( Oryza sativa L.MORPHOLOGY OF THE RICE PLANT ceae) . with round. sativa is indigenous to Asia. Seedling Morphology The grains of rice varieties that lack dormancy germinate immediately upon ripening. In dormant varieties. As the plant grows. O. Its color varies from colorless. fine branched roots form from the higher nodes on the. rather flat. sativa L. a rest period precedes germination and special procedures such as heat treatment (50°C. the coleoptile emerges ahead of the coleorhiza. The rice plant may be characterized as an annual grass. O. glaberrima is strictly an annual. 5 . A branch of the plant bearing the culm. while others may be diageotropic. possessing rootlets and root hairs.3 to . sativa mainly in a lack of secondary branching on the primary branches of the panicle and in minor differences related to pubescence on the lemmas and length of the ligule. In floating varieties. Roots The roots are fibrous. tall to floating varieties more than 7 m. leaves. tall. pale green to green. and auricles (Figs. O. hollow. The great majority of commercial varieties range from 1 to 2 m. roots and often a panicle is a tiller. sativa is a diploid species with 24 chromosomes. Under favorable conditions. jointed culms. The coleoptile later ruptures at the apex and the first seedling (primary) leaf emerges. days) or mechanical dehulling are needed to break dormancy in freshly harvested samples. and a terminal panicle. for 4-5.) beT longs to the tribe Oryzeae under the sub-family Pooideae in the grass family Gramineae (Poa- pears and is followed by two or more secondary seminal roots. The coleorhiza enveloping the radicle protrudes first if germination occurs in an aerated environment such as a well-drained soil. ligule. The second leaf that follows is differentiated into sheath. emerges as a tapered cylinder. Some of the adventitious roots are positively geotropic. in height. (2n =24). Seminal roots are later replaced by the secondary system of adventitious roots. The secondary adventitious roots are produced from the underground nodes of the young culms and are freely branched. The elongation of the mesocotyl elevates the coleoptile above the ground. The vegetative organs consist of roots. rice is adapted to an aquatic habitat. glaberrima Steud. sativa under series Sativa in section Sativae. Vegetative Organs The rice plant varies in size from dwarf mutants only . Its genomic formula is AA. O. glaberrima differs from O. coarse adventitious prop roots often form in whorls from the nodes above ground level. The primary seminal root (radicle) 1 breaks through the coleorhiza shortly after the latter ap1 Term in parenthesis indicates a synonym. the plant may grow more than one year. and leaves. 'or pale purple to purple. If the grain is submerged in water. 1. 1 and 2).long culm below the water surface. Biosystematists recently divided the genus Oryza into several sections and placed O. While the ensuing description is based on the ubiquitous O. the morphologic terms can also apply to the cultivated species of Africa. sessile leaf blades.

The nodal septum and internode may be differentially pigmented. The node bears a leaf and a bud. The septum inside the node separates two adjoining internodes. 2. generally increasing from the lower internodes to the Adventitious roots arise in both nodes and internodes and are usually found in the earlier formed ones. The bud the axil between the nodal septum 6 . The internodes of a culm vary in length. is made of nodes and internodes. Adventitious roots appear in the axis at the base of the internode.and the base of the sheath pulvinus. and glabrous on the outer surface. called a culm. Culm The jointed up of a series (nodal region) is inserted in stem of rice. The mature internode is hollow. The bud may give rise to a tiller. finely grooved.

The first rudimentary leaf at the base of the main culm is a bladeless. the enlarged. This node generally does not bear a leaf or a dormant bud but may give rise to the first 1-4 panicle branches. The latter give rise to tertiary tillers. The sheath pulvinus. 3). 3. The flower consists of six stamens and a pistil. The leaf consists of the sheath and blade. Floral Organs The floral organs are modified shoots. ligule and collar on the same plant may be differentially pigmented. Spikelets The spikelet is borne on the pedicel which is morphologically a peduncle. The nearly solid node between the uppermost internode of the culm and the axis of the panicle is the panicle base. 2. The extent to which the panicle and a portion of the uppermost internode extend beyond the flag leaf sheath determines the exsertion of the panicle. 1. The panicle base often appears as a ciliate ring and is used as a dividing point in measuring culm length and panicle length. and in the weight and density (number of spikelets per unit of length) of the panicle. The primary tillers originate from the lowermost nodes and give rise to secondary tillers (Fig. The flag leaf generally differs from the others in shape. The margins of the prophyllum clasp the young tiller with its back against the parent culm (Fig. the panicle (Fig. size. Varieties differ in blade length. one at each node (Fig. The primary branches may be arranged in a single or paired fashion. and angle. 2-keeled bract. form. The spikelets are pediceled on the branched panicle. shape. Auricles are small. Leaves The leaves are borne on the culm in two ranks. the dorsal. Varieties differ -in degree of exsertion. The main culm bears the largest number of leaves. Thus. Stapf and other systematists (cf. A spikelet is the unit of the inflorescence. color. the rachilla and the floret. angle. ventral and lateral parts of the collar may slightly differ in color. extending from the panicle base to the apex. The panicle axis (rachis) is the main axis of the inflorescence. with the perianth represented by the lodicules. the lower ones being larger in diameter and thickness than the upper ones. and pubescence. Varieties also differ in leaf number. The blades are generally flat and sessile. It is continuous and hollow except at the nodes where the panicle branches are borne. The leaf number on a tiller decreases progressively with the rise in tillering order. The swellings in the axils of the panicle where the branches are borne are the panicle pulvini. shape. 3). area. The lower internodes at the base of the culm are short and thickened into a solid section. When pigmented. The region about the panicle base is often called the neck. auricles. The junction of the sheath and blade is the collar or junctura. The secondary branches bear the pediceled spikelets. The upper surface of the blade has many ridges formed by the parallel veins. The leaf sheath is continuous with the blade. paired. The uppermost leaf below the panicle is the flag leaf. and tightness. The spikelet consists of the two sterile lemmas. The apex of the pedicel below the sterile lemmas is expanded into a lobed facet of varying size. It envelops the culm above the node in varying length. The prophyllum is also present between each secondary tiller and its tertiary tiller. glabrous or ciliate ligule. color. At the junction of the blade and sheath on the inside is a membranous. The sheath pulvinus is usually above the nodal septum and is frequently mistermed the node. The ligule varies in length. Tillers arise from the main culm in an alternate pattern. Panicle The panicle is borne on the uppermost internode of the culm which is often mistermed a peduncle. The collar often appears as a raised region on the back of the leaf. Varieties differ greatly in the length. ear-like appendages borne on either side of the base of the blade. The most prominent ridge on the lower surface is the midrib. and margin. A swelling at the base of the leaf sheath just above the point of its insertion on the culm is the sheath pulvinus. width. long) is considered as elongated. and angle of the primary branches. the prophyllum. two of which were reduced in development. A visually detectable internode (more than 5 mm. The panicle has a racemose mode of branching in which each node on the main axis gives rise to the primary branches and each of which in turn bears the secondary branches. cup-like apex is 7 . and shape from variety to variety. The terminal shoot of a rice plant is a determinate inflorescence. shape. The internodes also vary in cross-sectional dimension. 4). palea and the enclosed flower. Chevalier 1937) considered the spikelet of Oryza as comprising three fiowers.upper ones. The auricles may not persist on older leaves. A floret includes the lemma. 3).

3. Parts of a primary tiller and its secondary tiller. 8 .Fig.

Fig. 4. 9 . Component parte of a panicle (partly shown in this illustration).

and the included flower form the floret. the upper one generally being larger.Fig. The awn is a filiform extension of the keel of the lemma. outer glumes. empty glumes. 3-nerved palea. palea. homologous with a pair of true glumes and may be termed the rudimentary glumes. The middle nerve or keel may be ciliate or smooth. In some varieties. being always sterile.” “empty glumes. “empty glumes.. The apiculi may be separated into lemmal apiculus and paleal apiculus. 5).” the length of the latter. The extended tips of the lemma and the palea are the apiculi. and non-flowering glumes should be placed within inverted commas. 5-nerved bract which partly envelops the smaller. The lemma is the larger.g. The lemma. The surface of the lemma and the palea may be pubescent or glabrous. are the sterile lemmas (“glumes. During natural shattering or the threshing process. Therefore. such terms as glumes. A spikelet consists of a minute axis (rachilla) on which a single floret is borne in the axils of 2-ranked bracts (Fig. Parts of a spikelet. The sterile lemmas are generally shorter than the lemma and palea. a pair of lateral nerves on each side of the central nerve of the lemma may fuse to form a knob-like mucro on either side of the lemmal apiculus. The sterile lemmas may be equal or unequal in size. the pair of bracts above the rudimentary glumes should be designated as the sterile lemmas. The bracts of the lower pair on the rachilla. seldom exceeding one-third 2As Stapf’s interpretation is followed. e. the spikelet is separated from the pedicel at the junction of the lower sterile lemma and the 10 . indurate (hardened). The upper bracts or the flowering glumes consists of the lemma (fertile lemma) and palea.” “outer glumes”)2. 5.

The endosperm is enclosed by the aleurone layer which lies beneath the tegmen. The embryo contains the embryonic leaves (plumule) and the embryonic primary root (radicle). corresponding to the larger size of the lemma than that of the palea. The grain is the ripened ovary. The dehulled rice grain (caryopsis) is called brown rice because of the brownish pericarp. The base of the lower sterile lemma as is disarticulates from the pedicel is horizontal or oblique in appearance. Red rice owes its trade name to the red pericarp and/ or the red tegmen. Next in addition to amylopectin and stains dark blue to the pericarp are two layers of cells representing with potassium iodide-iodine solution. Anther dehiscence may coincide with the opening of the lemmas. if present.facet (rudimentary glumes) . rachilla. crude fiber. The hilum the embryo lies) are called white bellies. The starchy the remains of the inner integuments. transparent. or seed coat (often mistermed the testa). 6. A white is a dot adjacent to the embryo marking the point chalky region extending to the edge of the ventral of attachment of the caryopsis to the palea. sterile lemmas. 3. Some varieties are threshed by the fracture of the pedicel rather than by disarticulation.side and. these form the embryonic axis. 11 . The coleoptile is surrounded by the scutellum and the epiblast. and inorganic matter. The remaining part of endosperm. fats. firmly adhered to it (Fig. The nous) types. An. or immediately precede or follow it. with the lemma. Flower The flower proper consists of the stamens and pistil. Soft textured. Fig. forming a seed-like grain. and the awn whenever present constitute the hull or husk. the starch fraction contains amylose pericarp is fibrous and varies in thickness. non-waxy (non-glutiThe caryopsis is enveloped by the pericarp. In the waxy (glutinous) varieties. They represent the reduced perianth (calyx and corolla). the lodicules become turgid and thrust the lemma and palea apart. The pistil contains one ovule. The plumule is enclosed by the coleoptile and the radicle ensheathed by the coleorhiza. The lemma and palea and their associated structures such as the sterile lemmas. fleshy structures located at the base of the flower adnate to the palea. 6). The embryonic axis is bounded on the inner side by the scutellum (cotyledon) which lies next to the endosperm. The lodicules are two wale-like. At anthesis. The tip of the caryopsis is somewhat oblique. The short style bears the bifurcate. Chalky white spots often appear in the starchy The embryo lies on the ventral side of the spikelet next to the lemma. the starch fraction is composed almost entirely of amylopectin and stains reddish-brown with weak potassium iodide-iodine solution. The six stamens are composed of 2-celled anthers borne on slender filaments. rachilla. 1959). The surface of the caryopsis has ridges which correspond to those of the lemma and palea. toward the center of the endosperm is other scar at the tip of the caryopsis marks the called a white core. allowing the elongating stamens to emerge above or outside the open floret. Structure of a grain (adapted from Grist. In the common. The white starchy endosperm consists of starch granules embedded in a proteinaceous matrix. The lemma and palea close after the pollen grains are shed from the anther sacs. the tegmen endosperm also contains sugars. The relative degree of development of the abscission layer between the sterile lemmas and tile facet is reported to be associated with the ease of shedding. and the awn. palea. white spots occurring in the caryopsis is the endosperm which provides the middle part on the ventral side (side on which nourishment to the germinating embryo. plumose stigma. The rice fruit is a caryopsis in which the single seed is fused with the wall of the ripened ovary (pericarp). the vascular trace which is fused with the lateral parts of the scutellum. A long white streak on the dorsal side is called the white back base of the style.

P. often called the booting stage.dehiscing anthers in the terminal spikelets on the ish. The reproductive phase may begin before the maximum tiller number is reached. The bulk of the starch endosperm remains as the total milled rice (polished rice). and N. The nonrice and a small percentage of milled rice heavily 3The fortified with thiamin. Tanaka. the coarse outer bran layers. The members of the committee are R.proposed by a committee appointed during the Symposium colored riboflavin is added to the enriching agents. February 23-28.S. big brokens or second heads (broken kernels equal to or greater than half the length of a whole kernel). meiosis is usually . This is followed by the pre-tillering stage during which seminal and lateral roots and the first few leaves develop while the contents of the endosperm are absorbed by the growing seedling. Panicle development continues and the young panicle primordium becomes visible to the naked eye in a few days as a hyaline structure 1-2 mm. and Parboiled rice is rough rice which has been sub. Parboiling increases the percentage of As the grains ripen. The increase in the size of the young panicle and its upward extension inside the upper leaf sheaths are detectable as a bulge in the rapidly elongating culm. Yamada. to raise the vitamin and iron content slightly above unpublished nomenclature of growth stages of the rice plant the level present in brown rice. talc. screenings (broken pieces about 1/4 to 1/2 the length of a whole kernel). chips (small chips or particles of a kernel which can pass through a sieve having perforations of less than 1. trade terms.occurring in the microsporocytes (pollen mother cells) and macrosporocytes of the panicle. 2 percent pol. the rela. tegmen. 12 .47 mm. Enriched rice is a blend containing ordinary milled and turn yellowish in an ascending order. Growth Stages of the Rice Plant3 The vegetative phase of the rice plant begins with grain germination which is signified by the emergence of the radicle or coleoptile from the germinating embryo. The development of the fertilized egg and greatly among rice varieties of any particular endosperm becomes visible a few days following fertilization. mixture to make the finished product appear white. second heads (broken kernels at least half as long as a whole one). white pigments such as calcium oxide. medium brokens (broken pieces between 1/2 and 1/4 the length of a whole kernel). process. and iron phosphate terms used were largely adapted from a tentative. Grain development is a continuous geographic region. rice trade. after which some tillers die and the tiller number declines and then levels off. milling. total milled rice is separated into head rice (whole kernels). In U. long with a fuzzed tip. T. or 0. Owen. the hull is first removed from the grain (trade name: rough rice or paddy) in a sheller. Actually.055 inch). the embryo (germ) and small bits of endosperm constitute the rice bran. or about the period of the highest tillering activity. The visible elongation of lower internodes may begin considerably earlier than the reproductive phase or at about the same time. This phase is marked by the initiation of the panicle primordium of microscopic dimensions in the main culm. In the U. Matsushima. 8 percent bran.) and split kernels (pieces caused by a longitudinal splitting of the kernel). but agronomic terms such as the milk stage. on the Mineral Nutrition of the Rice Plant held at The International Rice Research Institute. S. titanium dioxide are also included in the enriching Chiu. and 1964. The increase in tiller number continues as a sigmoid curve until the maximum tiller number is reached.S. hard dough stage. Anthesis or Rough rice in the United States yields about blooming begins with the protrusion of the first 20 percent hulls.fully ripe stage are often used to describe the difjected to a steam or hot water treatment prior to ferent stages. niacin. Pollination and fertilization foltive proportions of the above components vary low. and 70 percent milled rice. A. S. The developing spikelets then become distinguishable. When the auricles of the flag leaf are directly opposite the auricles of the next lower leaf. small brokens (broken pieces which are smaller than 1/4 of a kernel but do not pass a sieve with perforations of 1. The tillering stage starts with the appearance of the first tiller from the axillary bud in one of the lowermost nodes.Trade Terms During the process of milling and polishing. The rice polish (white bran) refers to the inner layers of the bran removed during polishing. embryo. The pericarp.panicle branches. or thereafter. Evatt.4 mm. commonly called heading. When the yellow. the leaves become senescent head rice and the vitamin content of milled rice. Best. F. The corresponding trade terms proposed by the Food and Agriculture Organization of the United Nations (FAO) are: whole rice or head rice (whole or nearly whole kernels). N. aleurone layer. soft dough stage. and a small portion of the starchy endosperm are then removed as the bran. This is followed by panicle emergence from the flag leaf sheath. C. and brewers' rice (broken pieces which can pass through a 5½/64inch sieve).

point of spikelet separation). the zygote. HILUM. in some varieties. rough rice. BLADE (syn. eye). HULL (syn. It has no vascular tissue. The ripened ovary and its associated structures such as the lemma. COLEOPTILE (syn. containing oil and protein but no starch. sheathing leaf. brown rice). including the lemma. Structural changes (dissolution) in the abscission layer precede separation. APICULUS. husked rice. The round. In rice. germ. haulm). neck. 20. COLLAR (syn. 13. or tertiary. arista. ALEURONE LAYER. palea. such as a leaf or fruit. . junctura. sterile lemmas. 3. The cylinder-like. radicle. 14. 19. 5. The collar usually differs in color from the leaf sheath and blade. The joint between the leaf sheath and balde. lamina). rachilla. 15. if they survive the threshing 10. It is easily detached and removed in the milling process as pert of the bran. width. secondary. 11 . EPIBLAST. the culms and upper leaves may remain green when the grains are fully ripe. protective covering ma ana palea. new tillers may grow from the stubble of the harvested plants. six stamens. A unit of the spikelet.the lemma or palea. flat. consisting of hollow internodes jointed by solid nodes. 9. seed). sessile and free portion of the leaf. the awn if present. and the enclosed flower. The mature fruit of grasses in which the seed coat firmly adheres to the pericarp. CARYOPSIS (syn. adventitious roots and axillary buds. The embryo lies on the ventral side of the caryopsis next to the lemma. The caryopsis or dehulled grain. It persists sterile lemmas. ear-like appendages borne at the base of the blade and usually arising at the sides where the ligule and the base of the collar are joined. It is continuous with the leaf sheath. EMBRYO (syn. 13 . 4. 6. panicle. FLOWER. sickles). chaff). and the awn if present. that encloses the young plumule. considered to be a rudimentary cotyledon. 8.. which upon germination gives rise to a young seedling. paddy. BROWN RICE (syn. respectively. The embryo is appressed to the endosperm by the scutellum. the thickness of the abscission layer between the spikelet and the pedicel is reported to be associated with the ease of shedding in certain varieties. The sheath covering the process. and angle of insertion. 18. FLORET. Includes the lemleaf"). A filiform extension of varying lengths from the keel (middle nerve) of the lemma. derived from the fertilization of two polar nuclei in the embryo sac by one sperm nucleus from the pollen tube. stem. ENDOSPERM. depending on tillering order. and broken only for a short time after germination. consisting of the aleurone layer and the starchy endosperm. segment of the pedice1 are usually associated with the hull. 12. 2. AURICLES (syn.functioning leaves and culm tissues are termed dead straw. Structures such as the rachilla. beard). Sterile or under-developed ovaries enveloped by a well-developed lemma and palea should be termed empty or under-developed spikelets. Nutritive tissues of the ripened ovary. caryopsis. The two lodicules. Layer of cells related to the separation of a plant part. The miniature plant developed from the fertilized (diploid) egg. The plumule enclosed by the coleoptile and the radicle ensheathed by the coleorhiza form the embryonic axis in the embryo. Glossary of Morphologic Terms 1. The two apiculi may be distinguished as lemmal apiculus and paleal apiculus. and the pistil. . The extending tip of . The second and subsequent harvests from the crop are called the first and second ratoon crops. 7. It bears the leaves. The linear-lanceolate. ABSCISSION LAYER (syn. The peripheral layer of the endosperm. from the plant. A small structure opposite the scutellum in the embryo. smooth-surfaced ascending axis of the shoot. The endosperm is triploid. CULM (syn. husk. point of disarticulation. GRAIN (syn. 17. "first 21. Sometimes . COLEORHIZA. 16. it may be primary. A pair of small. Under favorable growth conditions. Blades on the same plant differ in length. palea. This structure may not persist on the older leaves. The basic parts of a mature embryo are the embryonic axis and the scutellum. EMBRYONIC AXIS. AWN (syn. leaf cushion). cargo rice). A scar on the caryopsis indicating the point of attachment to the palea.

extending from the base to the apex. It is light brown. more noticeable during panicle emergence. 30. The indurate. palea inferior. or adventitious roots. rhachis). 24. 25. rhachilla. originating from a node and enclosing the internode above it and sometimes the leaf sheaths and blades of the succeeding internodes. LEAF SHEATH (syn. sheath. LODICULES. The two other nerves are close to the margins. Its length can be measured only when the seedlings are grown in the dark or from the underground portion of the seedling. The smooth. and the sperm nuclei achieve the double fertilization process. non-glutinous rice). head. penetrates into the embryo sac. rudimentary glumes (facet). membranous structure seated on the inside of the collar at ita base where the blade joins the leaf sheath. between two successives nodes. The non-waxy type cooks drier than the gluey waxy type. 3-nerved bract of the floret which fits closely to the lemma. 23. basal portion of the pistil containing one ovule. A small bulge on either side of the lemmal apiculus formed by the fusion of the two lateral nerves. valve). “second leaf”). POLLEN GRAINS. 39. A small.” “glume. It is enclosed by the coleoptile. vagina). consisting of a haploid tube nucleus and a haploid generative nucleus. They are microgametophytes. and panicle branches. mesocarp and endocarp.22. with a median bundle but with no strong midnerve on the back. The pericarp is derived from diploid maternal tissue. 2-keeled bract enclosed by the leaf sheath with the back against the parent culm and its margins clasping the young tiller. The minute. 43.” “outer glume. stalk supporting branch. “glume. The determinate inflorescence of rice with a racemose mode of branching. NODAL SEPTUM. from nodes on the panicle. The internode between the scutellar node and the coleoptile in the embryo. 27. 40. In the young seedling. From the axils of nodes on the culm may arise a leaf. MUCRO (syn. 44. ear). The embryonic leaves of the young plant in the embryo. 42. glandular process).” flowering glume. PANICLE (syn. valvule). 41. PROPHYLLUM (syn. The pericarp layers may be differentiated into epicarp. PANICLE PULVINUS. The first seediing leaf without a blade that emerges next to the coleoptile. a tiller. The bulbous. A diminutive axis between the rudimentary glumes. 29. short basally and long apically. pale. ciliate or glabrous. Upon germination. NODE. flowering glume. The lower part of the leaf. peduncle). PANICLE AXIS (syn. The nearly solid node between the uppermost internode of the culm and the main axis of the panicle. solid (when young) or hollow (when mature) part of the culm. spheroidal structures (spores) in the anthers of a floret. They represent the rudiments of the perianth (calyx and corolla). 35. LEMMA (syn. Starchy endosperm in which the starch fraction contains both amylose and amylopectin. the branches or spikelets. A swelling in the axils of the primary panicle branches. The two scalelike structures which are adnate to the base of the palea. 31. The distinctly grooved. mesocotyl is the internode between the coleoptile node and the point of union of the culm and root. The a spikelet on the panicle end appears as a lobed cup. NON-WAXY ENDOSPERM (syn. panicle axis. It stains dark blue with weak potassium iodide-iodine solution. common. prophyll.” lower palea. 32. callus). inflorescence. The distal representing two foot stalk. ferti1e lemma. the pollen tube containing a tube nucleus and two sperm nuclei (gametes) grows down through the style. MESOCOTYL. PALEA (syn. PRIMARY LEAF (syn. It is similar to the lemma but narrower. 28. 5-nerved bract of the floret partly enclosing the palea. This node gives rise to the first primary branches of the panicle (1-4) and usually bears no leaf or dormant bud. PEDICEL (syn. 26. It is often bilobed. palet. 38. 37. INTERNODE. 34. RACHILLA (syn. upper palea. PANICLE BASE. upright. main axis of the panicle. The axis is hollow except at the regions (nodes) where the primary panicle branches are borne. 33. rachis. consisting of layers of cells which form a protective covering around the seed. The solid portion of the culm. The solid partition in the node separating two adjoining internodes. “inner glume. bearing pediceled spikelets and flowering from the apex downward. speckled reddish-brown. The wall of the ripened ovary. OVARY. coleoptyloid) . LIGULE. PLUMULE. palea superior. A thin. keeled. The indurate (hardened). 36. red or purple. 14 . PERICARP.

embryo and endosperm. The embryonic primary root ensheathed by the coleorhiza and the root cap? persisting only for a short time after germination. The portion of the embryo partly surrounding the embryonic axis. The term seed. seed coat). representing the inner cell layers of the inner integuments of the ovule. 47. 55. STARCHY ENDOSPERM. The true glumes of . typically including roots. The two sterile lemmas may differ in length and shape. 49. SEED. The bulk of the endosperm within the aleurone layer. WAXY ENDOSPERM (syn. the rachills and the floret. containing oil and protein for the germinating plant.” first and second glumes. A unit of the rice inflorescence consisting of the two sterile lemmas. TEGMEN (syn. The tegmen is often mistermed as testa which is derived from the outer integuments of the ovule and which is destroyed before the caryopsis ripens. It serves as an absorbing organ to transfer nutrients from the endosperm to the young seedling. The primary tillers give rise to secondary tillers and the latter to tertiary tillers. comprising roots and adventitious arise from the lower 52. A small swelling at the base of the leaf sheath just above its point of insertion on the culm. “glumes. sheath joint). SCUTELLUM (syn.” “empty glumes. Therefore. lower and upper empty lemmas). Waxy endosperm has an opaque appearance. vestigial glumes. 54. 46. 50. However. 45. The primary tillers originate from the lower nodes of the main culm. It is “glutinous” in character. opposite one another at the tip of the pedicel. 51. Starchy endosperm in which the starch fraction is composed almost entirely of amylopectin.a typical grass spikelet which are reduced to minute lobes in rice. actually refers to the grain.e. STERILE LEMMAS (syn. glumes. innovation). the seed is an inseparable part of the fruit (caryopsis). Rice pastries and high-quality rice wine are made from milled waxy rice. the waxy type of endosperm does not contain gluten. and fibers. The starchy endosperm also contains sugars. The two rudimentary glumes are considered to be a part of the spikelet. RADICLE. it becomes pasty and sticky when cooked. The two layers of cells lying next to the pericarp. fertilized egg including the seed coat. In rice. but which may or may not develop a panicle.” “outer glumes. TILLER (syn. The intravaginal vegetative branch of the rice plant. SHEATH PULVINUS (syn.. branch. The organs of age. 53. stool. sium iodide-iodine solution. The bulk of the starchy endosperm survives the milling and polishing process as the milled white rice. The two flowerless bracts at the base of the spikelet. the seed coat is firmly adhered to the maternal pericarp. cotyledon). absorption and anchorthe shoot. "non-flowering glumes.” lower and upper glumes. consistting largely of starch granules embedded in a proteinaceous matrix. culm and leaves. It rarely branches.the sterile lemmas. glutinous rice). SPIKELET. It stains reddish brown with weak potas-. as used in seeding or sowing. sterile glumes. 48. i. Often mistermed the node. and the fertile floret. facet). first and second glumes. 15 . fats. ROOTS. RUDIMENTARY GLUMES (syn. 56. The mature. All tillers arise in an alternate pattern. growing opposite the short-lived seminal secondary roots which nodes of the culm.

.

essential meteorological data. latitude.tillers have a pronounced spreading habit.g.the angle is intermediate between erect and spreading. such as height. grain shedding. Oñate 1964. 3. Leaf sheath : The sheath color of the lower leaves is readily classified.) is measured from the base of the plant to the tip of the longest leaf of seedlings pulled at random at a given date (14-21 days) following seeding.000 cultivated varieties. and others. growing period. Juvenile growth habit: Juvenile growth habit may be measured from the angle of the tillers observed from the entire plot prior to the maximum tillering stage. the distance (in cm.. Standard color charts for plant tissues are available from the Munsell Color Charts for Plant Tissues (1963). pathological. Some of the exotic or extreme types are described under “Mutant Traits” which follows. sativa L. red. e. Seedling height: For seedlings grown in the seedbed or directly sown in the field. physical measurements are more meaningful than generalized descriptive classification of tall and short. 1965. (1938) and Ghose et al. In such cases. b. particularly those designed for direct seeding. grain weight. Erect . Leaf color: a. Plants with prostrate habit in early growth may assume a less spreading form later in their lives. Some of these are being used to catalog The International Rice Research Institute’s collection of 10.an angle of 30° or less from the perpendicular. (1960). Information on sampling methods or sample sizes for quantitative traits may be obtained from Institute publications (IRRI 1964. they may be modified to suit local needs. maturity and size. vary greatly in growth habit. As some of the criteria are empirical measures. Economic traits of a physiological.BOTANICAL AND AGRONOMIC TRAITS USEFUL IN VARIETAL CLASSIFICATION AND IDENTIFICATION he thousands of cultivated varieties (cultivars) T of O. Oñate and Moomaw 1965). there is no precise definition or means of measuring seedling vigor. leaning more than 60” from the perpendicular. b. cultural methods. c. Prompt emergence and rapid growth are generally desired in commercial varieties. Description and color plates are given by Hutchinson et al. and intensity of pigmentation have shown marked variability when the plants are grown under different sets of environments. Spreading . Blades: Foliage color is readily distinguished in the seedling stage. 2. Although varieties differ greatly in seedling vigor and rate of growth. and soil fertility level. 17 . form. Experiments showed that certain characters which were believed to exhibit discontinuous variation such as awn length. Color classes are shades of green. it is desirable to indicate the environmental conditions under which the particular trait or traits were observed. Intermediate . a. or quantitative nature may be used to aid in varietal identification. grain dormancy. size and structure. The commonly used plant characteristics and the methods of recording them are enumerated below. For quantitative traits which continuously vary within a variety when grown under different environments. A strict botanical classi- Seedling Characteristics fication based on morphological differences does not provide sufficient criteria to embrace the enormous diversity in rice. and purple. Standards may be based on local varieties if such standards are indicated in the published results. early and late. Juvenile plants of the intermediate type are less prostrate than those with the spreading habit. 1.

(1960). purple stripes. several shades of purple. Additional tests for physiological characters of rice seedlings have been described by Oka (1958). medium (20-34 mm. Nagai 1958). Some floral organs may also lose their color or fade as the plant matures.) : the lowest elongated internode which is longer than 5 cm. and virus diseases (IRRI 1965. the bacterial leaf blight disease ( X. Length of mesocotyl. Standard color charts may be referred to for finer differentiations. Seedling reactions to specific chemicals: Varieties differ in their seedling reactions to specific chemicals. Collar: The collar may be colorless (white). b. most of the tropical indica varieties are susceptible to the phytotoxicity of organo-mercuric fungicides. 5. Angle : Two measurements can be taken on the same blade. 1. (1938).tip dips lower than the collar). length and density of pubescence. and Ghose Adult Plant Characteristics The characteristics of the adult plant are recorded during or shortly after anthesis.4. 7.. shades of purple and purple lines. Resistance to blast and other diseases: Seedling leaf reaction to the blast fungus (Piricularia oryzae) can be readily determined in a blast disease nursery. sp. Root color: Colorless (white) or red (under sunlight). length. gold. and long (35-50 mm. coloration (colorless.). green. and 91° or more. 6. 5. Blade: The uppermost leaf below the flag leaf on the main culm is taken as a representative blade. Xanthomonas translucens f. Color: colorless (white). b. a.). or shades of purple). and dark green. 3. and color are recorded on the main culm at full blooming. Auricles: Auricles are examined for presence or absence. The color of the inside sheath base (sometimes called the leaf axil) just above the pulvinus is another taxonomic criterion. Length: short (5-19 mm. Area: Leaf area can be estimated by the product of length and width. blade largely straight and erect but curved near its tip). Varieties also differ markedly in seedling reaction to the bacterial streak disease. green or purple. The other is the angle of blade openness measured up to the apex of the blade. e. Operational details may be obtained from the procedure for establishing a Uniform Blast Nursery (Ou 1965) . blade generally long and its. f . Colors of the sheath on the outside are green. blade angle may be classified as erect (angle of attachment and angle of blade openness nearly equal. The differential effect of herbicides on rice varieties may also be used to differentiate varieties. It varies from colorless (white) to green and shades of purple. For instance. (1938). 6. Length ratios of primary leaf/coleoptile and mesocotyl/coleoptile may also be used to differentiate varieties (cf. horizontal .) from the junction of the blade and leaf sheath to the tip of the blade. in total darkness. 4. Leaf sheath: Sheath color is taken on the first leaf below the flag leaf. c. Ligule: Characteristics of the ligule are also recorded on the first leaf below the flag. Ramiah and Rao (1953). Ramiah and Rao (1953). Angle of flag leaf: Angle of 0-30' at full blooming is rated erect. Width: Measured at the widest portion of the blade. and shades of purple. c. descending. Color of internode surface: green. Other colors are full purple. 2. full purple. One is the angle of attachment of the blade measured near the collar. curving (angle of openness larger than that of attachment. Therefore. Pubescence : Pubescent or glabrous (including smooth surface and ciliate margins). 18 . 31-60°. intermediate: 6190°. coleoptile and primary leaf : Mesocotyl length is measured from 7-day old seedlings germinated at about 30°C. oryzicola ) . Culm: Characteristics such as outer diameter. a. straight blade). 1966). b. and purple lines. blade gently curving throughout its length) and drooping (angle of openness much larger than that of attachment. whereas the japonica varieties are largely resistant. Length: The distance (in cm. oryzae ( X. Outer diameter (in mm. colorless roots may turn reddish brown because of the deposit of ferric hydroxide on the root surface. Color plates have been given by Hutchinson et al. and Ghose et a1. Sheath color variations have been illustrated by Hutchinson et al. Mesocotyl and coleoptile development may serve as indices of seedling vigor during emergence. oryzae ). recurvate at the tip (angle of blade openness slightly larger than the angle of attachment. d. purple stripes. and purple wash of a spreading type (Ramiah and Rao 1953). 7. As roots grow older. purple margins. Pubescence of fringes: glabrous or ciliate.).. In descriptive terms. when recording pigmentation it is desirabIe to indicate whether the plant is at the blooming or mature stage. Color: Green is divided into pale green. a.

ochraceous-buff. d . seashell pink. Angle of primary branches: erect. straw. c. Length: long. Colors are light yellow. gold. June-November. Pubescence : glabrous or pubescent (indicate length and density). Apiculus (examined first during anthesis and then at maturity) : a. d. Plants with bending or buckled culms are called weak or lodged. b. Color plates have been given by Takahashi (1957). amaranth purple. Chang 1964b). light purple. or extra long (longer than lemma) . This trait is first rated following panicle emergence by gently pushing the tillers back and forth at a distance of about 30 cm. (1938). rose red. Detailed information on straw strength may be obtained by (i) mechanical culm breaking or pulling devices. Upright plants are considered sturdy. c. Color illustrations have been given by Hutchinson et al. from the ground. 13. russet. Form: equilateral (paired branching) or unilateral (one-sided branching). or a. faded pink. Nodal septum: The color of the septum is best seen by slitting longitudinally the lower portion of the culm and examining the cut surface. Additional observation at maturity is made to record the standing position of the plants. warm buff. b. Rachilla: comma-like. or full purple). Number: Culm or tiller count includes both panicle-bearing and non-bearing tillers. pansy purple. they are termed brittle. Pubescence : glabrous or pubescent . shades of purple. Lemma and palea: a. 10. (ii) a brass chain hanging from the panicle base to give cLr (lodging resistance factor) estimates. nearly equal or unequal (one-sided) in length. c. and blackish red purple. 11. short or long trichomes (indicate the length and density of trichomes and specific areas of measurement). brown spots (piebald). or tip awn. Panicle: May be cup-like. or intermediate. or partly stained. Color plates have been given by Takahashi (1957). IRRI 1964. compact. tawny (light to dark brown). 14. yellow. (iii) Terminally awned: short awns are present on spikelets near the tip of the panicle branches. shades of purple (including red). b. or sooty black. brown furrows. d. red. 16. Colors are colorless. Takahashi (1957). c. reddish brown. 8.5 percent aqueous phenol solution for 24 hours. pomegranate purple. Phenol staining: Grains are soaked in 1. purple. b. pink. gold. drained and air-dried. gold. elbow-like. Length: short (less than one-third of lemma). or intermediate. Ramiah and Rao (1953). or purple. drooping. Color at anthesis : straw white. Length: The distance in centimeters from the ground level to the panicle base. and faded purple. Strength : Culm strength varies among varieties and also within a variety with changing environments. c . brown. red or purple. long (more than one-third). short. Color at ripening: white. Color: colorless (white). 15. straw white.et al. (1960). Sterile lemmas (recorded as the terminal spikelets start maturing) : a. faded red purple. medium. and Ghose et al. straw. (i) Fully awned: all spikelets on the panicles are awned: but awns often vary in length. or black. or purple dots. 9. teristic. Hull color is then recorded : unstained. straw. e. purple spread. 12. Color: colorless (white). (iv) Awnless: awns are absent and do not develop under any condition. blackish brown furrows. The ratio of bearing tillers to the total number of tillers is also a varietal charac-. pale yellowish green. tawny (light to dark brown). Length: measured from the panicle base to the tip. shades of purple (purple tips. 19 . gold. or (iii) P/E estimates (ratio of critical straw strength to the modulus of elasticity) derived from culm measurements (cf. Awn (recorded as the terminal spikelets start maturing) a. Data are taken at the Institute during the main (wet) growing season. If culms break readily. Color at maturity: White. brown. b. (1960). tyrian rose. Color at anthesis: green. russet. Stigma color: This is examined during anthesis using a hand lens. (ii) Partly owned : awned and awnless spikelets are present on the same panicle. piebald and mottled patterns. and shades of purple. Sheath pulvinus: The pulvinus can be colorless (white). This bending test gives some indication of stiffness and resilience. green. Type: open. pink. entirely stained (dark brown). Presence .

The duration of grain development generally varies from 25 to 40 days. June-November. Another commonly used measure is the date of panicle emergence (heading). Presently. 19. h. full heading. (iii) Partly enclosed (panicle is partly enclosed by the flag leaf sheath). natural long-day conditions) with a duration comparable to or not more than 10 days longer than that grown under the 10-hour treatment (or comparable. Oka (1954). 60 percent emergence of all panicles. Consequently. of light) would differentiate most varieties. Guevarra and Chang 1965). Formulas for estimating photoperiod sensitivity have been given by Chandraratna (1966. Planting most of the photoperiod-sensitive varieties under short-day length would result in fewer tillers. For tropical varieties. Among photoperiod-sensitive genotypes (i) of diverse geographic origin. 116-130. The number and individual length of elongated internodes are characteristic of a certain variety under a given environment and are often associated with growth duration (cf. information on photoperiod sensitivity and/or thermosensitivity may be obtained. the growth duration of a photoperiod-sensitive variety at a given location or latitude is determined by the optimum photoperiod of the variety and the dates on which the critical photoperiod prevails at that latitude. The total growth duration of a rice variety generally may be resolved into 4 components: (a) basic vegetative phase. i . no standard methods are available to evaluate readily the photoperiod response or thermosensitivity of many varieties. the spikelets are evenly distributed along the primary or secondary branches. different planting dates usually affect the total growth duration. Therefore.) several times over panicle samples placed on a flat or inclined board and counting the percentage of dropped grains. 161-176. and (d) reproductive phase from panicle initiation to maturity. Exsertion : Exserted (panicle base is clearly above the flag leaf sheath). Ratings are: (i) Tight. and 16 hr. middle heading: and more than 90 percent emergence of all panicles. g . Internode elongation pattern: Plant height is largely the summation of elongated internodes (including the panicle axis) on the culm. natural shortday conditions). or longer. f. Shattering can also be determined in the laboratory by rolling a weighted cylinder (about 1 kg. and earlier maturity. The effect of thermosensitivity on growth duration is generally less than that of photoperiod sensitivity. intermediate. Clustering: in most varieties. (ii) Intermediate: 25-50 percent of grains removed. (c) thermosensitive phase. 101-115. (b) photoperiodsensitive phase. two controlled photoperiods (10 hr. Weight (of panicles on the main culm dried to 13 percent moisture content of grains). shorter plants. Density (number of spikelets per unit length of panicle) : dense. especially in the tropics. 176-190. Days from sowing to 5 percent emergence of all panicles in a plot may be called days to first heading. and intermediate. When photoperiod chambers are available. some varieties have two or more spikelets clustered on the’ panicle branches at irregular intervals. Shattering: Tight. The degree of clustering varies from 2 to as many as 48 on a secondary branch. 146-160.: few or no grains removed. of which 4 to 9 are elongated 5 cm. 191-205. or shattering. if duplicate plantings are made during different seasons. 18. varieties often differ in the optimum photoperiod at which the panicleinitiation process is critically affected. and Katayama (1964).e. Occasionally. 17. (ii) Partly exserted (panicle base appears at the same level as the top of the flag leaf sheath). Shattering can be measured in the field by gently grasping by hand the mature panicle and applying a slight rolling pressure. 131-145. However. (iv) Enclosed (panicle is entirely enclosed by the flag leaf sheath). Maturity: Maturity is computed in days from seeding to ripening of more than 80 percent of the grains on the panicle. lax. Internode elongation patterns represented by ideograms or internode length/culm length ratios for specific internodes facilitate comparison of varieties under identical or different treatments (IRRI 20 . (iii) Shattering: more than 50 percent of grains removed. plant height and other quantitative traits are measured in the main (wet) season. Plant height: Measured on the main culm (or the tallest tiller) at or following anthesis from ground level to the tip of the panicle. Varieties showing a difference of 10-20 days in heading date may be classified as weakly photoperiod-sensitive: those with a difference longer than 20 days are definitely photoperiodsensitive. Rice varieties generally have 12 to 22 internodes. 1966). the following maturity ranges are applicable: 100 or less. At the Institute. and 206 or more. A photoperiod-insensitive variety would initiate panicles under the 16-hour photoperiod (or comparable. Varieties also differ in the duration from anthesis to full maturity.

9 mm. shape and weight: a) Length. Dead : leaves become non-functional even before the grains are fully mature.) : longitudinal dimension measured from 10 well-developed grains as the distance from the base of the lowermost sterile lemma to the tip (apiculus) of the lemma or pales. Percentage of fertility more than 90 75-90 50-74 10-49 less than 10 Classification highly fertile fertile partly sterile sterile highly sterile 22.) : lateral diameter measured from 10 grains as the largest distance between' the two lateral sides in the middle part of the caryopsis. 24.51 mm.50 mm. 7). length is measured to a point comparable to the tip of the apiculm (Fig. It may be expressed as a ratio of photosynthetic leaves to the total number of leaves on the culm. b . Grain dimensions.0 mm.0 less than 2. Volumetric weight such as test weight or liter weight is another useful varietal characteristic. 7). 5. Hull percentage: Hulls are readily removed in a sheller or dehuller.0-2. useful to rice agronomists and breeders.1 2. 20.61-7. shape. In the case of awned varieties. d) Shape: generally expressed as a ratio. 1966 .4-3. between length and width. c) Thickness (mm. FAO scale for milled rice more than 7 mm. a. Spikilet or pollen fertility: The mean percentage of fertility is obtained from the percentage of welldeveloped spikelets (or pollen grains) in the panicle and sampled from a number of panicles in the plot.0-7. The complement of hull percentage is the percentage of brown rice. 6. 7.1964.0 2.classification schemes that have been suggested are as follows: Size (length) Extra long Long Medium or middling Short USDA scale for milled rice4 more than 7. 23. A photo-enlarger with a calibrated easel is used at the Institute for the above classification. Seven grain weight classes are given by FAO . Dimensions. 21. The proportion of hull to grain (rough rice) on a weight basis is another useful varietal characteristic. whichever is longer. 5.0 more than 3 2. 5. shape. Hull percentage may vary from 16 to 35 percent among varieties. Guevarra and Chang.0-6. and 100-kernel weight of brown or milled rice are additional criteria for identifying varieties.51-6. (mm. Two . Photosynthetic leaves at maturity: This characteristic is not commonly reported but may prove.60 mm. Length and width measures of the grain. Photosynthesis: leaves retain their green color when panicles ripen.60 mm. and weight of brown or milled rice: The size. FAO scale for Shape (length/ USDA scale width ratio) for milled rice brown ria Slender Medium Bold Round more than 3.) : dorsiventral diameter messured from 10 grains as the distance across the lemma and-the palea at the widest point (Fig. A screw micrometer or a dial-type vernier caliper is used. Fig. e) Weight: measured from 100 grains dried to 13 percent moisture. 21 . 1966).39 less than 2 4 Unofficial scale used by USDA rice researchers and is not the basis for official USDA marketing classes. b) Width (mm.1-3. 6. less than less than 6 mm.

culm rot. Protein content is determined by analytical methods. Known examples include rice blast. b) Gelatinization temperature : Gelatinization temperature indicates the temperature at which starch granules swell irreversibly in water with a simultaneous loss of birefringence. 30.) measured with an amylograph suggests certain parboiling and canning characteristics of rice samples (Halick and Kelly 1959). common : dark blue staining reaction with potassium iodide-iodine solution. light red. c) Pasting viscosity: The difference ("setback") between peak viscosity of hot-paste (95°C. reddish brown. rice stunt and other virus diseases. and white back. Environmental factors such as temperature. intermediate (70°74°). golden. Juliano et al. medium. Potassium iodide-iodine solution is prepared by dissolving 1 g. The differences in cooking and processing behavior are largely due to inherent differences in the chemical make-up of the starchy endosperm rather than to grain size and shape. grain size and shape are generally associated with certain cooking and processing characteristics. large (22 to 28 g. of iodine in 100 ml. stem maggots. Rhizoctonia seedling blight. Embryo size: Small. nitrogen supply. Weight: 3 weight classes are given by FAO for 1. Non-waxy or non-glutinous. milling and/or cooking. Whether a variety is scented or not may be detected during anthesis. Cercospora leaf spat. Shape: No standard measures are available for tropical varieties. 29. large. water. Waxy or glutinous: brownish staining reaction with weak potassium iodide-iodine solution. Pasting viscosity is affected by amylose content and protein content. indicating that only amylopectin is present in the starch granules. Varieties can be further differentiated in their reaction to specific pathogenic or physiologic races 22 . sheath blight. 26. Resistance to diseases and insects: Rice varieties differ in their reaction to a specific pathogen or insect pest. reddish purple.). e. Amylose content may be determined analytically (Williams et al. round. Amylopectin/amylose ratio in starchy endosperm : The amylopectin. Alkali-resistant samples indicate high gelatinization temperature. medium. e) Aroma: scented or non-scented. of potassium iodide and 0. the granule-swelling method./mm. Size: Whole kernels and broken pieces may be separated by a sizing device and expressed as a ratio or percentage. light. b. amylose ratio expresses the relative proportion of each type of starch in the endosperm. Commonly used designations are slender (fine). bold (coarse). and the loss of birefringence of starch under a polarizing microscope. red. indicating the presence of amylose in the starch granules. and intermediate. white-tip disease (nematodes). The identity of the volatile chemical (s) in scented rice has not been determined. Physical features of milled rice: a. c. d. opaque. and high (75°-80°). d) Protein content: The exact role of protein content in determining cooking quality is not well understood but it is known to be affected by environmental and nutritional conditions under which the crop is grown. grayish brown. Fusarium seedling blight. Three classes are generally recognized: low (62°-69°). 1958) or estimated by the starch-iodine-blue test (Halick and Keneaster 1956). Chemical properties of starchy endosperm related to cooking characteristics: Among rice varieties. 28.) and cooled paste (50°C. b. bacterial leaf blight.3 g. Pericarp and tegmen colors include white. and the stem borers. leaf-hoppers. Pericarp and tegmen color: The color of brown rice is determined by pigments in the pericarp and tegmen.2 f. Some of the inherent varietal quality differences are given be- low.25. and storage conditions also affect the cooking behavior. Amylose content and gelatinization temperature appear to be genetically independent (Beachell and Stansel 1963. Chalky spots: white belly. white core. and small (less than 22 g. There are many exceptions to this classification. Short-grain varieties are usually more cohesive and firm than long-grain varieties. Gelatinization temperature also may be estimated by the amylograph test. fluffy cooking features.) . and full purple (almost black). Helminthosporium leaf spot. 27. and flat. 1964). Most long-grain Varieties tend to become dry and fluffy when cooked and the cooked kernels do not split or stick together. Translucency : translucent. Medium-grain varieties have generally intermediate features.000-kernel weight of milled rice very large (more than 28 g. The starchy endosperm of all rice varieties is white. bacterial streak. a.) . brown. It may be estimated by soaking milled rice for 23 hours in a weak alkali solution at 30°C. The extent of endosperm disintegration gives an estimate of relative gelatinization temperature. a) Amylose content: High amylose content is associated with the dry. Hardness: determined by a grain hardness tester as g.

33.0 percent and still produce a fair yield of grain.5 percent and 1. and (6) maturity ranges of 100 to 150 days. These differences are crucial in sub-tropical and temperate regions where rice is sown early in the year. Irrigation water of low temperature readings early in the planting season has caused severe mortality to rice seedlings in California and Hokkaido. (3) low to medium tillering ability. and summer varieties on the basis of the harvest season. As mentioned before. Other classification schemes based on growing period are : main and off seasons . As a general rule. they can stand 2 to 3 meters of water without obvious adverse effects. nonfloating varieties and the fioating varieties. and (c) grain weight. No clearcut morphological or physiological criteria are yet available to differentiate rice varieties into lowland and upland types.e. There is also a group of varieties intermediate between the common. In other staple cereals. Rice varieties also differ in their reaction to certain physiological disturbances. i.. Ratooning ability: Rice varieties differ in ratooning ability following initial harvest. medium. Well known samples are specific varietal reactions to different races of the blast fungus. and wet and dry seasons. Yields from upland fields are generally lower than those from lowland fields. Yield and yield components: The three basic physical components of grain yield per unit area are (a) the number of panicles per unit area. Experiments have shown that most of the socalled upland tropical varieties gave higher yields when grown under irrigated conditions than under a natural. low air temperatures during the period between panicle development and pollen fertilization cause high sterility. simple and reliable . early. Reaction to physiological diseases. (4) non-sensitive or weakly sensitive to photoperiod. The term saline-resistant or saline-tolerant varieties has been used to denote varieties than can tolerate salinity levels between 0. 32. Seasonal adaptation : Rice varieties of India and Pakistan are divided into autumn. Floating varieties are generally described as long duration (160-240 days). although such terms are widely used as cultural designations. or is low-lying without adequate irrigation facilities. Other examples are physiological diseases such as “Akiochi” and “Akagare” caused by a deficiency in certain nutrient elements and/or an excess of harmful products of extreme soil reduction. (b) the number of welldeveloped grains per panicle. (2) vigorous seedling growth to compete with weed growth. and late seasons. rapidly rising water levels during the peak of the monsoon season permit only the growing of the floating va- rieties. Heritable differences among Japanese varieties have been obtained to facilitate selection for more tolerant strains. Cultural adaptation: Rice culture is generally divided into lowland and upland culture. haphazard type of intermittent flooding.tests for resistance to salinity need to be formulated. But attention must be given to uniformity in cultural factors such as water and nutrient supply when comparing varietal differences in ratooning ability. Lowland culture refers to continuous flooding of the fields except for occasional drainage. They are known to withstand 5 to 8 meters of standing water. spring. controlled irrigation. upland type of cropping in Japan and certain parts of west Africa to the rain-fed “upland’ ’fields of the tropics where the rice plant grows in flooded soil for a substantial portion of its life cycle. Varieties also differ markedly in tolerance to low air temperatures and strong air movement shortly after seedling emergence from the soil. photoperiodsensitive varieties with many internodes which can elongate rapidly to cope with rapid increases in water level. Some of the “upland” varieties yielded as high as or even higher than certain lowland varieties when grown under flooded conditions. Known examples are differences in varietal resistance to straight head which is largely caused by prolonged flooding of some soils. Designating a variety as upland is not necessarily related to the resistance to drought of the variety either in the seedling or the adult stage. 31 . varietal differentiation based on specific physiological characters is preferred to cultural designation. first and second seasons . Under comparable growth 23 . In many river deltas of tropical Asia. Called “deep-water’’ varieties. 35. A number of the so-called upland varieties grown in the tropics have the following characteristics in common: (1) rapid emergence from the soil following direct seeding. There is sufficient varietal diversity to enable rice breeders to select cold-water resistant strains. Again.of one pathogen. In the tropics an “upland” field means that either the field is elevated ground which gravity-fed irrigation system cannot reach and where water-retaining devices are not available. At high latitudes. the temperate zone japonica varieties are more tolerant than the tropical indicas. 34. pure isolates of the rust fungi are sometimes used as a tool in purifying a breeder’s seed of commercial varieties. Experiments have shown that the above divisions were based largely on photoperiod sensitivity and thermosensitivity. But upland culture embraces a continuous range from the strictly non-irrigated. winter.

the terms indica. each variety shows a fairly consistent composition of grain yield in terms of the three components. dark green leaves Short. many panicles per plant). hybrid sterility. and (c) an intermediate type. the above scheme of dividing rice varieties into geographic races is rapidly losing its significance. Varieties differ in the duration and intensity of dormancy (Jennings and de Jesus 1964). tropical areas. Leaf characters: Varieties differ in leaf dimensions. 1964). Pakistan. and seedling reaction to organo-mercuric compounds. Japanese workers (Kato et al. and serological reaction.C.S. Other criteria related to leaf area index and useful in differentiating varieties are the light transmission rate (I/I0) and the extinction coefficient (K). physiologicaI characters of germinating seedlings and adult plants. resistance of seedlings to KClO3 toxicity. Philippines. 1928) divided cultivated rice into two subspecies.. c. b. In 1928-1930. phenol staining of hulls. plant and grain morphology. Rice varieties have been divided into three groups on the above basis : (a) “panicleweight” or “heavy-eared” type (large and heavy panicles. Varieties which fall into the japonica group have been isolated in semi-wild conditions from Nepal. and Taiwan varieties developed in recent years. “indica” and “japonica. plot yield is more meaningful than single-plant yield. density. whereas the japonica group (= ‘kêng’) consisted of varieties from northern and eastern China. Oka 1958. IRRI 1964. roundish grains Medium tillering Short plant stature Awnless to long awned Dense and long hairs on lemma and palea Low shattering Hard plant tissues Varying sensitivity to photoperiod JAVANICA Broad.” on the basis of geographical distribution. and other. Geographic designation : Since two hundred B. The indica group (= ‘hsien’) included varieties from Ceylon. Despite these shortcomings. Hayashi and Ito 1962. India. stiff. This is particularly true with U. and the phenomenon of intervarietal hybrid sterility was much more complicated than a simple classification of three groups. Taiwan. Morinaga 1954. Other tests of potential value: In addition to the above methods and criteria. few panicles per plant). Ceylon.” to designate the bulu and gundil varieties of Indonesia. thick grains Low tillering Tall plant stature Awnless or long awned Long hairs on lemma and palea Low shattering Hard plant tissues Low sensitivity to photoperiod Other criteria used by Japaneee workers to classify varieties are endosperm characteristics. (b) “paniclenumber” or “many-eared” type (small and light panicles. Therefore. Japan and Korea. Later studies with larger collections of varieties showed that the morphological and physiological variations among the three geographic groups were largely continuous. and glutinous. Among the three. d. rice varieties of China were recorded under three groups: ‘hsien’. Morinaga and Kuriyama 1958) later added a third group. “javanica. The leaf area index (LAI) of the blades is highly characteristic of a variety when grown under a given set of environmental condition8 and is useful in comparing the efficiency of leaves in utilizing sunlight. the light transmission rate is the most readily measurable criterion (cf. southern and central China.conditions. light green leaves Broad. somewhat flat grains Profuse tillering Tall plant stature Mostly awnless Thin and short hairs on lemma and palea Easy shattering Soft plant tissues Varying sensitivity to photoperiod JAPONICA Narrow. the Jeypore Tract of Orissa State in India. Japanese workers (Matsuo 1952. Java. Tanaka et al. air-dried to about 14 percent moisture. Grain yielding ability is also used as a varietal characteristic. ‘kêng’. and northern Thailand. and javanica are often used by rice scientists in Asia as convenient designations to indicate different plant and grain types. The grain/straw weight ratio is another criterion which has shown relatively high consistency in certain varieties and may therefore be used as an additional basis for varietal differentiation. Grain yield data supplemented by data on the three components provide more information for comparative or diagnostic studies. Spodogram analysis: The shape. several others which show promise in differentiating varieties are cited below : a. Grain (seed) dormancy: Germination tests of freshly harvested panicles. 37. will give an indication of dormancy. 36. Numerical symbolization of pigmentation in plant parts : In view of the complexity of the anthocyanin distribution among plant parte and the various properties of color expres- 24 . japonica. density and arrangement. Hybridization on a wide genetic basis has further confused the classification scheme. lightgreen leaves Slender. The above three groups and their general morphological and physiological features are summarized as follows : INDICA Broad. When used as such. and distribution of silica cells in the epidermal tissues of blades and leaf sheath are highly characteristic of certain varieties.

the arrangement 25 . Grist 1959. the nature of the starchy endosperm. Beale further subdivided each group into seven classes on the basis of pigments in the stigma. Sativa The great majority of the classification schemes found in the literature emphasize varietal differences of a regional scope (cf. was given on as many as 78 items for each variety. However. apiculus. a centralized agency is needed to work toward planting. In addition to anthocyanin pigmentation and grain dimensions. Classification of Cultivated Varieties of O. Chandraratna 1964) and therefore have limited application.1 issues. value. the information on individual varieties is based on data furnished by breeders in the variety’s home country. leaf axil. and the blade. recording and cataloging existing commercial varieties for morphological and agronomic characteristics under a uniform set of conditions and to preserve the valuable germ plasm when the existing stock of varietal diversity rapidly dwindles. f. Consequently. Protein fractions of plant tissues: Studies at the Institute have shown that certain indica and japonica varieties differ in the protein fractions of leaf tissues (IRRI 1964). This has helped rice workers select and exchange experimental materials. a variety of multivariateanalysis techniques are now available to assist in evaluating similarities between taxonomic units and the ordering of these units into groups on that basis. value and chroma in 5 selected plant parts. But the task of continuously developing identification and classification schemes for commercially important varieties of a country or region should be largely the responsibility of national governments concerned. and chroma). international and interagency cooperation is needed to pool all available information on cultivated varieties in a form that would be readily accessible to all interested workers. 1964). the Food and Agriculture Organization of the United Nations has published a World Catalogue of Genetic Stocks (Rice). The 35 combinations supposedly embraced the total varietal diversity in Burmese rice. of spikelets on the panicle branches. leaf sheath. Therefore. in which available informatior. Ito and Akihama (1962) suggested the use of a numerical scale to combine recordings of the hue. and the form of spikelet separation from the pedicel were the criteria used in other classification schemes. While none of the presently available classification schemes is entirely satisfactory for the thousands of rice varieties under cultivation and probably none will ever serve the needs of all interested workers. using the Munsell color charts.sion (hue. Other aspects of chemical plant taxonomy have been discussed in a symposium on the subject (Swain 1963). the size and shape of the sterile lemmas. An extension of this scheme may facilitate the card cataloging of varieties. This is illustrated by Beale’s (1927) classification of rice varieties from lower Burma in which five major groups were recognized on the basis of grain length and the length/width ratio. They also indicated that the greenness of leaves is correlated with nitrogen responsiveness and chlorophyll content (Tanaka et al. Statistical approach to varietal classification: When a number of taxonomic units of either a quantitative or codable nature are recorded. the presence or absence of awns. since it is impossible for one agency to develop a catalog that would contain all of the information desired by various workers and which would also apply to all of the varieties when grown at different locations. totaling 1. Principles and techniques have been given by Sokal and Sneath (1963). the pubescence of the lemma and palea. The world collection of the Institute is being investigated and maintained in the above manner to meet this need. e. some of the quantitative data have limited applicability when the variety concerned is grown under a markedly different set of environmental conditions. supplemented by similar measurements of the dehulled grain. This is particularly true with varieties of the temperate zone when grown in tropical areas under cultural practices adapted to the profuse tillering tropical varieties. Since 1950.

.

when used as the first letter of a gene symbol. and I-Pla. 6 P. e. coleoptile. (1960). and Wh for white hull. awn. Ghatge and Subrahmanyan (1960). Genetic information on other mutants of a more exotic nature have been given by Jones (1933). Some of the P genes also have several alleles. green. ex. Red pericarp and tegmen (red rice) is controlled by two complementary genes. The genes controlling anthocyanin pigmentation are basically three complementary genes: C5. hull (lemma and palea). Through the interaction of the above genes. Dwarf plants form discrete classes in segregating generations and are characterized 27 . Variations in Non-Anthocyanin Pigmentation The coloration of plant parts such as gold. sterile lemmas. Thus. nodal septum. leaf axil (inner sheath base). and P controls the distribution or localization of anthocyanin in specific plant organ or organs. and stigma.g. Anon. pink. Pl. Nagai (1958). Non-anthocyanin pigmentation generally involves a single pair or a series of alleles. Chang and Jodon 1963). and Ghose et al. The mogen. Inhibitor genes (I-) are known to suppress the effect of the distribution genes. and several shades of purple. Takahashi (1957). Pin for internode. brown. sheath pulvinus. The P and Pl alleles also have a pleiotropic effect upon the pigmentation of the other plant organs. The color variations are colorless (white). a great variety of color expressions and intensities are observed in nature.. Ramiah and Rao (1953). The more commonly observed ones are reviewed under eight convenient groupings. A. Rc and Rd. leaf tip and margin. Jodon (1957). A wide array of mutant traits has been reported in rice. and Chandraratna (1964). Modifications in Size and Shape A common modification in size is dwarf stature which is about one-third to one-half the height of normal plants. gh for gold hull. C is the basic gene for producing chro5 Letters in italics are IRC-recommended gene symbols (cf. 1963. 1959. tegmen. Recent tabulations on gene symbols. I-Pl. Bh genes for black hull. Pau for auricles. Nagao (1951). blade. Ps for stigma. pericarp. Psh for leaf sheath. genetic ratios and authors are given in two IRC reports (Anon. A controls the conversion of chromagen into anthocyanin. mutant trait is defined as a variant that is relaA tively discrete from the normal or parental type and is inherited in a simple Mendelian manner. Pl for leaf blade. such as Bf alleles for brown furrows on hull at maturity. Variations in Anthocyanin Pigmentation The plant parts which may be pigmented with anthocyanin are the apiculus. ligule. Inhibitor genes such as I-Bf and I-H have been reported. the color of the caryopsis is speckled reddish-brown.MUTANT TRAITS normal type refers to that of the great majority of cultivated varieties.. and Px for leaf axil. Colors for the above have been designated by Hutchinson et al. exceptions: Ph. internode. C and A are genes with multiple allelic series. Definitions of specific mutant traits and assignment of the IRC-recommended gene symbols are given in the attached glossary. and P6. Examples are P for apiculus. When Rc alone is present. red. I-P. denotes anthocyanin color in a certain plant organ or organs. Prp for pericarp. collar (junctura). the mutant traits are variants of a more extreme nature than those commonly observed in a small group of commercial varieties. Ramiah and Rao (1953). Plw and Plt. and sooty black does not involve anthocyanin. Chang and Jodon 1963). (1938). leaf sheath. auricles. Pi. Ghose. Anon. 1963. midrib. H alleles for dark brown furrows on hull at blooming.

bend node (bn) . . Differences in plant stature between the tall tropical varieties and these short varieties are controlled by a partially dominant allele and a few modifying genes (Chang et d. viridis and xantha. recessive genes (d1. Dw2). and other physiological characters mentioned in preceding sections also involve modifications in the chemical composition of the plant organ or tissues. lazy or ageotropic growth (la). lazy (la). ligule. lax panicle (dn). or Gm for long). spreading panicle branches (spr) . d2. collar. growth habit. translucency and chalkiness of the starchy endosperm (wb. Physiological diseases showing dark brown or blackish mottled discoloration of leaf blade are inherited as single recessive characters (bl). White streaks or stripes (ws) may also be found on the blade. sinuous neck (sn). Although some of the above traits. belong to this class. . non-floating growth (Dw1. variations in maturity or photoperiod sensitivity. Modifications in Growth Habit Some of the well-known modifications in growth habit are erect (er) vs. and pubescence generally involves differences in one allele. zebra stripe (z). twisted leaf (tl) .. non-exserted panicle (ex). Other Modifications in Structure Marked variations in the structural features of plant organs include rolled leaf (rl). 1947) has described a number of chlorophyll mutations in barley. Presence or Absence of Structures The presence or absence of awns.. cleistogamous spikelets (cls) . xantha (I-y) . These single-recessive dwarfs have no economic value in a breeding program. Pigmentation. maternal inheritance of chloroplasts has been indicated. Modifications in Other Physiological Characters Other well-known modifications in physiological characters involve a complex of chlorophyll deficiencies. pale yellow (y). green and white stripes (fs or gw). Chlorophyll deficiencies occur in a variety of expressions. . spreading. erect vs. wc) . and differences in ratooning ability. variations in grain dormancy. ) of a pleiotropic nature control the dwarfed growth. and tip-burn yellow (tb) are lethal types and are detectable immediately following seedling emergence Chlorina (chl) . auricles. culm. and panicle length. and collar is generally inherited as a unit (Lg vs. undulate rachis on the lower panicle branches (Ur). Color plates for the above types have been given by Ramiah and Rao (1953). claw-shaped spikelets (clw) . and spikelets persisting throughout the growing period. In others cases. Lg for liguled. Sk2. nitrogen-responsive and high yielding indica varieties from Taiwan. sterile lemma length (g. Taichung (Native) 1. leaf sheath. anti Gl for pubescence. and resistance to specific diseases and insects. an inhibitor for tall plant height (I-T) may be involved. and virescent (v) are non-lethal and the seedlings usually regain the green color later in growth. were described as simple Mendelian characters they are probably polygenic in inheritance. chlorophyll deficiencies. presence of a certain structure is generally controlled by the dominant allele. and phenol staining of hulls (Ph). Other simply inherited differences in size and shape involve grain length( lk alleles. open hull (o). e.). blade width (nal for narrow blade). extra lemma (lmx) . A number of independent. lg) in a pleiotropic manner. I-geo-tze and Dee-geo-woo-gen. striata. waxy endosperm (wx). fragrant flower (fgr). and notched or twisted kernel (nk or tk). floating vs. The presence or absence of the ligule. such as panicle density and shattering. The short (circa 100 em. clustering of spikelets on the panicle branches (Cl) . . 1965). Gustafsson (1942. such as An for awned. poly-embryonic grain (me). Modifications in Chemical Composition Some of the simple modifications in chemical constitution of plant parts are extremely brittle culm and leaves (bc). In other cases. Albino (al) . The above chlorophyll deficiencies are largely controlled in each case by a single recessive allele. albina. lutescent (lu) .by under-sized grains and proportionally thickened plant parts. scented endosperm (Sk1. beaked hull (Bd). auricles. The. viz. grain shape (Rk for roundness). shattering (Sh or th). ). neckleaf. verticillate rachis (ri). 28 .g. glossy leaves. which also may apply to rice. gametic and zygotic sterility. Plants of intermediate (sub-normal) height with normal panicles and grains may be called short or semi-dwarf plants. These differences in growth habit also are known to be controlled by specific chemicals (hormones or growth substances). for long). leaf discolorations. depressed palea (Dp). triangular hull (tl-i) . double awn (da). Some of the above variations in size and shape are probably more of a quantitative nature. multiple pistil (mp).

Seedlings are yellowish green as a result of chlorophyll deficiency. Nagai (1958). Sterility occurs in the reproductive organs at different . and Chang and Jodon (1963). chlorina (chl) .Sooty black color of the hull is controlled by two or more complementary genes which are not related to genes for anthocyanin production (C. Interpretations were offered either in terms of a single dominant gene for resistance (Pi. short awn (an1 An2 An3 or an1 An2 an3). Grain dormancy has been reported as a dominant trait controlled by two complementary genes (Sd1. due to low content of a-cellulose in cell walls.are dark brown. Non-dehiscent anther is a common type of male sterility. Rice workers are urged to look for additional mutant traits of economic value. 5. Sd2) or several genes. Plants are often smaller in stature than normal. The seedlings die shortly after emergence. Ce) or a dominant gene for susceptibiity (or inhibitor gene). Differences in growth duration have been reported to be controlled by a single gene. albino (al) . particularly at maturity. P). Chromosome aberrations such as deletions. a study of maturity by the component-analysis approach may clarify this complex trait.Physiologic leaf spots are non-pathogenic physiologic diseases showing dark brown or blackish mottled discoloration of leaf sheaths and blades. haploidy. 8.types are blotches of white spots on the blade and discolored spots on the hull (hsp).Spikelets show a clumped arrangement on the primary or se- 29 .Auricles are reduced to protuberances with 1 to 2 cilia or without cilia. and other malformations. rudimentary auricles (au) . 3. clustered spikelets (Cl) . asynapsis (as) and desynapsis (ds) which occur during meiosis are cytologically detectable.The furrows on the lemma and palea. Morphological steriles include empty anther sacs. Among physiological disturbances. ) which showed cumulative action in conditioning resistance to different pathogenic races. A. The inheritance of dormancy is probably more complicated than generally held as the hull. or tip awn (an1 an2 An3). awned (An) . 11. opaque endosperm with high lysine content. In other cases. 13. Intervarietal hybrid sterility of a genic and/or chromosomal nature is often observed in progenies of distant crosses.Leaf blade has white spots which give a pie-bald appearance. Glossary of Mutant Traits and Gene Symbols 1.The awned condition is classified according to length as long or full awn (An1 An2 An3 or An1 An2 an3). and male sterility.The culm forms an angle at the node. Resistance to drought and low air temperature have been reported to be under polygenic control.The tip of the lemma is recurved over the palea. replacement of stigma by supernumerary stamens. brittle culm (bc) . polygenic inheritance would gain the attention it deserves. Previous studies on the genetics of the rice plant are marked by a predominance of major genes and duplicate loci. Anon. similar to a twisted stem. As discussed in the preceding section. brown furrow (Bf) . 9. leaves. and each may play a specific role in conditioning dormancy. Exceptions to the above generalized patterns of inheritance are included in reviews by Ramiah and Rao (1953). . (1963). Resistance to fungus diseases such as blast and Cercospora leaf spot was studied genetically either with pure isolates or composite cultures of the pathogen. aerial branching at nodes (Br) . Awnless condition has the formula an1 an2 an3. 2.The culm. Details are given in recent reviews by Chandraratna (1964) and Chang (1964). bent node (bn) . 4. and panicles are extremely brittle. blotched white . 6. As the genetical studies of economic. solid or semi-solid culms. Pi2.Young seedlings are white due to the absence or degeneration of chloroplasts. beaked hull (Bd) .New shoots and roots develop at nodes high on the culm following the cutting of panicles. 7. quantitative traits receive more attention by rice breeders and geneticists. 12. translocations. Ef alleles for earliness. blackhull (Bh) . or Lf alleles for lateness. inversions. triploidy. photoperiod sensitivity ( Se alleles) was interpreted as the basic factor in late maturity. including yellow endosperm.stages of development either as morphological aberrants or physiological disturbances of the reproductive process. and embryo are involved. tetraploidy. Similarly. medium awn (An1 an2 An3 or An1 an2 an3). . varietal reactions to insect pests were reported in terms of single allelic differences. physiologic leaf spots (bl) . replacement of pistils and stamens by glumes. and aneuploidy also cause sterility. pericarp. 10. The above mutant traits found in some of the other staple cereals have been utilized in breeding programs. or duplicate genes for resistance (Pi1.

polycaryoptic (mp) . few hairs may be found on the margins of the blade.The panicle is enclosed by a spathe-like leaf or leaf sheath arising at the panicle base. synonymous to “parrot beak.A small wedge-shaped depression occurs at the middle portion of the abdominal side of the brown rice. The plants are often characterized by undersized grains and proportionally thickened plant parts.” 16.Long and dense hairs on the leaf blade. . depressed palea (Dp) . extra lemma (lmx) . 39. The blade stands upright at base. 24. 32. hull colors (Hm. 21. 22.The hulls show a golden yel- low color at maturity. lax panicle (Lx) . Plants also are short. 19.Long. erect growth is dominant to lazy. Hi.The culms are erect and closely grouped. fine stripe (fs) . The awns may have unequal length. dense or compact panicle (Dn)-Avery dense arrangement of spikelets on the panicle branches.Panicles with few lateral branches and sparsely distributed spikelets on the branches are called lax. additional bracts may arise at the base of the panicle branches. is present. 26. probably synonymous with shaggy hull. synonymous with juncturaless. 37. Goldhull is recessive to straw colored hull.Spikelets have an extra glume between the fertile lemma and the sterile lemma.Hairs arc scarce or absent on the hull and/or the leaf blade. 28.The spikelets have undersized paleas which are overlapped by recurving lemmas. with compact panicles and small spikelets.condary panicle branches with two or more spikelets per branch. 30 . 34. claw-shaped spikelets (clw) .Seedlings are normal green at first.Dwarf plants are about onethird to one-half the height of normal plants. erect growth habit (er) . lazy (la) . resulting in a large number of spikelets per unit length of panicle. 14. 18. The exserted type is dominant over the enclosed condition which is due to a shortened internode below the panicle base. liguleless (lg)–Leaves lack collar.The sterile lemmas (“outer glumes”) exceed one-third the length of the lemma and palea.Several pistils with multiple ovaries are present in the spikelet. Gm. Erect growth is recessive to the spreading or procumbent habit which has culms growing obliquely outward. long glume (g) .Green and white stripes on the leaf blade and occasionally on the lemma and palea. Lax panicle has also been used to denote open and spreading forms.Leaf blades have a glossy surface to which water easily adheres in large droplets. some extreme spreading forms have been mistermed lazy. owing to shortened internodes and/or fewer elongated internodes. lutescent (lu) . very hairy (Lh) .Seedlings show fine greenish white longitudinal stripes at tip and margin of leaf blade. fuzzy hull (Fg or Lh) . The spikelets therefore cannot open at anthesis. exserted panicle (Ex) . 27. 40. ligule. In crosses with ageotropic lazy (la) type. dominant to ordinary pubescence.Plants grow prostrate rather than upright because of an ageotropic growth habit. hullspot (hsp) . but the chlorophyll gradually disappears and the plant ultimately dies. double awn (da) . glabrous (gl) . and auricles. cleistogamous spikelets (cls)-Thelemma and palea unite at the lower end. Hf) . 31. notched kernel (nk) . Stripes often disappear in the tillering stage. 20.Small tissue areas on the lemma and palea which are thin and transparent and appear discolored. dense hairs on the hull result in a fuzzy appearance. In rice literature. Ho. Bunched arrangement of spikelets in groups of 10 to 48 is termed superclustering. Gh. Dwarfs form discrete classes in segregating populations. glossy . smooth. dwarfs (d) . Another dominant gene.A number of hull colors are non-anthocyanin colors which appear on the lemma and palea only when the straw hull gene.The panicle emerges fully from the flag leaf sheath at panicle emergence. 33. 23. polyembryonic (me) . The two sterile lemmas may have unequal length. Occasionally. 30. neckleaf (nl) . 29. green-and-white striped (gw) . 38. 15. 36. controls the sterile lemmas which are longer than the lemma and palea.The palea is underdeveloped and shows a depressed appearance. synonymous with polyhusk. 17.Spikelets have awns developing on both lemma and palea. 35. In a glatbrous strain.Multiple embryos are present in one caryopsis. 25. goldhull (gh) .

For color illustrations. 46. 42.A chalky white spot occurs in the middle part of the caryopsis on the ventral side (on which the embryo lies). 60. 61. opposed to straight neck. Rc and Rd. 52.The long caryopsis shows a slight twist in the middle portion. but this condition is non-lethal. opaque appearance. forming a half cylinder. 56. Rc.The chalky white.The spikelet appears triangular because the lemma is so shaped.Brown. The leaf apexes wither and die. 69. gray. 53.The lemma and palea cannot close after they have opened at blooming time. sometimes as a result of underdeveloped midrib or absence of midrib in the blade.The portion of the uppermost internode of the culm below the panicle base is curved and wavy. 55. 50 spreading panicle branches (spr) . speckled reddishbrown tegmen and pericarp.Longitudinal white streaks appear in the leaf shath. The starch in the white belly is soft. localizers or distributors (P-). The starch is non-waxy. chromogene ( C alleles). red pericarp and tegmen. and white pericarp and tegmen colors were reported. refer to Nagao (1961) and Ramiah and Rao (1953). This condition is caused by mechanical compression exerted on the branches owing to the premature extension of the axes while they are inside the flag leaf sheath. shattering (Sh or th) . waxy endosperm (wx) .Chalky white hull is dominant over straw-colored hull and epistatic to goldhull (gh) . 63. golden. zebra stripe (z) . 47. 44. 64. 48.The primary panicle branches extend obliquely outward so that they appear spreading and lax. and spikelet throughout the growing period.Young seedling is nearly white or slightly yellow as a result of the delayed development of chloroplasts. scented kernel (Sk) . (1938). undulate rachis (Ur) .The starchy endosperm is soft and crumbly with a dull. short non-dwarf plant (I-T) . Purple pericarp is controlled by Prp or PI alleles. opaque endosperm .Yellowish seedlings die after the second foliage leaf is formed.41. 46. white hull (Wh) . red.Seedings have pale yellow leaves. soft portion extends to the edge of the ventral side and toward the center of the caryopsis. stains reddish brown with weak potassium iodide-iodine solution instead of the blue black color of the common or nonwaxy type of starch. 62. 43. The basic genes involved are activator (A alleles). xantha (l-y) . white belly (wb) . purple. 58. 54. especially the lower ones.Seedlings have pale yellow leaves and perish shortly after germination. inhibitors (I-).Grains contain aromatic substances in the endosperm which give a strong aroma when the polished rice is cooked. 51.Seedling leaves have transverse. and modifiers. 31 .Panicle branches are arranged as a whorl around the basal node of the panicle axis. Later the plant gradually turns green. virescent ( u ) .Shattering or easy threshing are partly correlated with the degree of development of the abscission layer between the spikelet and the facet of the pedicel.The primary panicle branches. refer to Hutchinson et al. rolled leaf (rl) . long twisted kernel (tk) . white stripe (ws) . yellow leaf (y) . culm. verticillate rachis (ri) . and Chose. the panicle and grain sizes are about normal. in which the starch fraction is composed of nearly 100 percent amylopectin. 49. However. alternating chlorotic bands which later disappear.Leaf blade margins are incurved.Plant height is shorter than normal. tipburn yellow (tb) . white core (wb2) . have undulating secondary axes. blade. triangular hull (tri) . 57. sinuous neck (sn) . open hull (o) . Ghatge and Subrahmanyan (1960). pericarp and tegmen colors . Ramiah and Rao (1963). For details. pigmentation of vegetative parts .Waxy or glutinous type of starchy endosperm. Takahashi (1957).Anthocyanin colors in vegetative organs are controlled by the complementary C-A-P system.

Ramiah. and wild taxa is supported by a grant from the United States National Science Foundation (GB-2417). Seetheraman. Nagamatsu. E. S. Shastry. and Joao de Carvalho e Vasconcellos have commented on the glossary of morphologic terms and related items. Bor. Prior to the preparation of the manuscript. The authors also wish to thank C. and B. Thomas K. Jodon. Y. R. SI. S. E. Beachell. C. Hsieh. S. T. The work of recording and cataloging the Institute's ten thousand cultivated varieties. Roy Adair. N. R.ACKNOWLEDGMENTS This compilation includes the contributions of many workers who are interested in bringing uniformity to the taxonomic and genetic nomenclature of rice. Akira Tanaka. Ramiah. Rogelio Feliciano. R. E. Hiroshi Ito. the two glossaries were circulated among a number of agrostologists and rice researchers. genetic testers. Colin McClung. Soderstrom. H. Nelson E. Parthasarathy. Roy Adair. Jennings. H. Juliano. and Man-emon Takahashi offered suggestions on the mutant traits. Jodon. I. Nelson E. T. K. A. Sampath. S. X. Terrell. V. Mao. Tuguo Tateoka. K. Hubbard. S. Oka. O. Man-emon Takahashi. C. Vergara who critically reviewed the manuscript. P. C. B. Seetharaman. E. 32 .

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Ghose. London. Inst. 1963. H. Bull. Sci. Jodon. Moomaw. pl. Rao. 11. Oñate. 12(4) : 18-29. 39. International Rice Research Institute. 35.. Annual Report. Genetics anu Breeding of Rice. Chemical Plant Taxonomy. W. B. K. I. Principles of Numerical Taxonomy. Kadam. China 51(N. Pal. M. Internatl. Tsai. F. E. Chang. J.360 pp. H. Chandraratna. A. I. 3: 1-80. Takahashi. Dept. Hiroko Morishima. Gustafsson. Rice Comn:Newlet. Anonymous. Philippine Agr. 1: 1-96. III. T. B. T. N. Y. 441-446. 38. V.S. 1960. 1951.) 47 : 771-782. 19. Variability in rice. Garcia. Internatl. Facul. Rice Breeding and Genetics. 3. Agr. R. New Delhi. B. Advn. and E. analysis of plant height. Genetic symbols for rice recommended by the International Rice Commission. 5) : 592-616. 543 pp. 3) : 266-362. Agr. Res. E. Longmans. The description of crop-plant characters and their ranges of variation. Inst.. M. Plant Breeding 18: 79-89 33. Laguna. J. J. 1947... B. Agr. 8(pt. Assoc. 2nd ed. Genet. 6 pl. 2. 1933. 1964. 1964. Monitoring of gene symbols in rice. pl. T. N. Inst. Varietal resistance as a method for rice stem borer control. Newslet. and V. Sethi. E. 10. and M. S. J. L. 1964. and H. Takahashi. Rice Comn. Laguna. Indian Council Agr. Mitra. Ghatge. pp. II. 1965. ARS 34-28: 1-56. Rao. 1965. A.. I-VI.) : 1-8. B. J. and B. Rice gene symbolization and linkage groups. I. A. Wu. Newslet. and M. 1964. Res. 474 pp. A. 1942. K. S. 50: 266-362. 8. 360 pp. A. and N. Tagumpay. S. ICAR. Genet. 15. Res. 37. 17.. 7. M. 1963. T. 1958. The plastid development in various types of chlorophyll mutations. Monogr. maturity and other quantitative traits in the cross of Peta x I-geotze. 49: 1-8. J. Thompson... Internatl. Part IV. S. ICAR. Tech. B. Internatl. London. T. Academic Press. Sneath. Nagai. US. Res. Nagao.. Oka. 50(pt. San Francisco. 335 pp. M. Los Baños. Anonymous.. Ou.. (US. 4: 181-212. J. 1959. Virginia R. Growth habit of rice plant in the tropics and its effect on nitrogen response. Mutations in agricultural plants. S. 8(4) : 1-7. Mimeo. Gustafsson. Indian J. 6: 47-48. Philippines. P. Sokal. New Delhi. M.. Rice Res. B. S. Faculty' Agr. Agr.. Hereditas 33: 1-100. Intervarietal variation and classification of cultivated rice. Monog. 1958. 843 pp. 14. Tateno. Tech. 1957. J. 1938. Statistics in Rice Research. 1964. O. A proposal for an international program of research on the rice blast disease. ed. Ramiah. 1963. Indian Council Agr. Internatl. Tanaka. 5. Inheritance of some of the more striking characters in rice. 1965. Rice Breeding and Genetics. Analysis of apiculus color genes essential to anthocyanin coloration in rice. Williams.32. Hereditas 28 : 483-492. Hutchinson. Navasero. Indian J. T. XXXVIII-LIII. Ramiah. 4. Rice Res. 13. Huang. 1963. 13(1): 15-21. T. Rice Res. New Delhi. Jodon. Rice in India. T. 1958. T. Agr. 1953. Oñate. A. F. Food Chem. M. Present knowledge of rice genetics and cytogenetics. Internatl. 9. and J. Parao. Sci.. Bates. T. Res. Baltimore. Japonica Rice: Its Breeding and Culture. V. R. Chang.. Bull. 250 pp. Freeman. H. 48: 181-192. Genetic . Pathak. 1957. Los Baños. Jones. 34. Sci. Chang. S. B. The Rice Blast Disease. R. Ser. Rice Comn. 19. 40. 1964. Yokendo. Agr. Hokkaido Univ. Variability of characteristics in rice research. M. Tokyo. N. Genic analysis and linkage relationship of characters in rice. K. Subrahmanyan. Mutant Traits and Gene Symbols 1. W. L. T. Thadani. 35 . Analysis on apiculus color genes essential to anthocyanin coloration in rice. Johns Hopkins. 1953. Indian Council Agr. 1957. C.D. 12. 6. Hokkaido Univ. Varietal differences in the amylose content of rice starch. K. Philippines. K. Inheritance of characters in rice. Swain. H. V. 36. Hered. 359 pp. C. 41. Alam. 389 pp. Rice Res. 1965. C. Inst. Internatl. and P. R. Ramirez. M. Ramiah. 16. 42. G. and K.

25. 22. 29 Double awn. 15. 13. 7. 19 strength. 18 length. 21. 24. 28. 28. 18. 12. 5 Chromosome aberrations. 13. 11. 13. 7. 18. 27. 13. 14. 19. 25. 5. 17. 11. 15 Collar. 5. 19. 11. 21. 29 Brown furrows. 13. 12 outer. 23 Albina. 14 Disarticulation. 29 Blade (Also see Leaf). 13 pubescence. 13. 25. 17. 13. 30 shape. 11.SUBJECT INDEX Abscission layer. 29 Coleoptile. 20 Beaked hull. 27 Dominant gene. 18. 29 Black hull. 29 Diploid. 15. 11. 18 area. 13 inner. 10. 29 Dwarfs. 27. 11. 15 Empty lemmas (See Sterile lemmas) Endosperm. 28. 28 color. 27 length. 28. 12. 13. 7 number. 11. 20 main. 13. 28. 27. 23 seasonal. 27 absence or presence. 28. 18. 29 Brown rice. 6. 28. 22 Bud. 19. 29. 25. 22 Embryonic axis. 29 36 . 19. 28 color. 13. 11. 18 diameters. 30. 27. 12. 18. 6. 30. 22 Asynapsis. 24. 6. 14. 27. 31 Adaptation. 7. 21. 29 size. 7 width. 11. 13 Cargo rice (See Brown rice) Caryopsis (Also see Brown rice). 11 Disease resistance. 13. 10. 14. 18. 29 Awn. 30. 12 Branch (See Tiller) Branching at nodes. 27. 21. 7. 10. 10. 29. 18. 15. 14. 30 Cleistogamous spikelets. 28. 18. 28. 29 Anthers. 30 Clustered spikelets (Also see Panicle). 19 length. 27 color. 31 Chaff (See Hull) Chlorina. 31 angle. 5 length. 31 Chromogene. 12. 7. 13. 20. 28. 13. 27. 15 Alleles. 6. 31 Chromosomes. 11. 13. 19. 18 color. 13. 29 Anthesis. 29 Chlorophyll deficiencies. 5. 12. 28 Albino. 30 Ear (See Panicle) Egg. 18 pubescence. 27. 27 color. 29 Brittle culm. 30 Duplicate genes. 7. 22-23 Distribution gene. 21 shape. 29 Claw-shaped spikelets. 13. 31 Activator gene. 29 Bracts. 18. 28. 29. 19. 11. 7. 21 percentage. 12. 15. 18 Color charts and plates. 29 Beard (See Awn) Bent node. 11. 18. 13. 30 absence. 27 lemmal. 28. 7. 14 axillary. 25 Dense panicle (Also see Panicle density). 12. 13. 12. 13. 17. cultural. 30 Anther dehiscence. 31 Complementary genes. 30 color. 31 color. 10. 28. 5. 12. 30 Desynapsis. 14 paleal. 29. 13. 27. 11. 5. 30 Depressed palea. 15 Embryo. 28. 18 rudimentary. 12. 27. 18. 19. 12. 18. 7. 11. 19 Cultivars (See Cultivated varieties) Cultivated varieties. 28 Aleurone layer. 14. 31 Apiculus. 7. 17. 7. 29 Basic vegetative phase. 20. 28 Blooming (See Anthesis) Blotched white. 18 Coleorhiza. 29 Auricles. 28. 13. 28. 14 Bran. 31 Cotyledon (See Scutellum) Culms. 27. 29 Anther sacs. 13. 14. 13. 28. 19 Arista (See Awn) Aroma. 5. 12. 12 Anthocyanin pigmentation.

24 Geotropism ageotropic. 25. 15 “sterile”. 15. 10. 12. 5. 30 diageotropic. 20 Height. 31. 12. 17. 14 Filaments. 5. 12. 17. 12. 15 first and second. 13 Exotic types (See Mutant traits) Exserted panicle (Also see Panicle exsertion). 7. 22. 15 flowering. 30 juvenile. 12 germination. 30 maximum tillering. 28 Floral organs. 10. 30 Husk (See Hull) Husked rice (See Brown rice) Indica. 22. 22. 15. 5. 11. 20. 28. 15 Flower. 14. plant. 21. 25 Foot stalk (See Pedicel) Fragrant flower. 25. 12 panicle development. 21. 15 “first and second”. 31 seedling. 25 world. 13. 28. 30 rudimentary. 12. 15. 12 Glabrous. 28. 28 waxy (glutinous). 15 Inhibitor genes. 11. 30 Growth duration. 13. 20. 31 starchy. 7. 20. 20 panicle initiation. 18. 20. 11 Fine stripe. 17. 12. 14. 30 “First leaf” (See Coleoptile) Flag leaf. 10. 30 Glumes. 23. 15 vestigial. 15 International Rice Commission (IRC). 21 Hullspot. 14. 14. 15. 5. 27 International Rice Research Institute (IRRI). 30 37 . 17 lazy. 11. 11.. 12 booting. 14 inner. 21 weight. 12. 11. 17. 28 erect. 13. 12. 10. 11. 28 Insect resistance. 14 “lower and upper”.14. 28. 6. 7. 24.30 spreading. 28. 15 “non-flowering”. 20 middle. 27.Endosperm (continued) common (non-waxy). 12 internode elongation. 5 Geographic races. 25 Internodes. 30 Growth stages. 29 germination. 30 percentage. 14. 15 Glutinous rice. 7. 7. 28 Fruit (See Caryopsis) Fuzzy hull. 31 dormancy. 31 Grain. 11 Floret. 23 Integuments inner. 11. 14. 13. 12. 11. 12. 28 thickness. 9. 12. 15 outer. 14 Gelatinization temperature. 28. 24 Green-and-white-striped. 27. 20 pollination and fertilization. 14. 11. 17. 20 full. 18. 19. 21 Grain/straw ratio. 12 length. 29 annual vs. 28. 30. 24 Inflorescence (Also see Panicle). 17. perennial. 31 Enriched rice. 23 panicle emergence. 12. 14. 28 Grain (continued) shape. 5 components of. 7. 5 geotropic. 30. 30. 18 sheath. 22. 17. 12. 24 width. 12 Epiblast. 3. 15 “outer”. 11. 12 meiosis. 10. 12 vegetative. 10. 10. 13 Hull. 15 “empty”. 19. 28. 12 grain development.collection of cultivars. 17 Hilum. 25 “Indica” subspecies. 5 Fertile lemma (See Lemma) Fertilization. 20 effect of photoperiod. 12 Haulm (See Culm) Head (See Panicle) Heading (Also see Panicle emergence). 25. 12 reproductive. 28. 30 Extra lemma. 21. 21. 10. 19. 20 first. 30 Glossy. 15. 13 Food and Agriculture Organization (FAO). 12 tillering. 31 pre-tillering. 13. 5 Genus. 28. 22 Genomic formula. 27 recommended gene symbols. 11. 15 “glume”. 3. 14 “glumes”. 12. 5 Germ (See Embryo). 15 true. 27. 24. 13. 17. 30 Gametes. 15 Goldhull. 23. 29 colors. 20 effect of planting dates. 22. Floating habit (See Growth habit). 30 Eye (See Embryo) Facet (See Rudimentary glumes) Family. 20 Growth habit.

30 Leaf. 7. 7 photosynthetic. 27 blade (Also see Blade) . 15 Paddy (Also see Grain). 12 Maternal inheritance. 19 secondary. 18. 20. 7. 11 Notched kernel. 20. 20 initiation. 19. 10. 7. 20 length. 30. 21 chalky spots. 31 primary. 12. 21. 30. 13. 29 color. 19. 7. 21. 13. 18 color. 24 “Javanica” subspecies. 24 Junctura (See Collar) Keel. 29 Panicle pulvinus. 15. 17. 3. 21. 20. 28. 28. 30 color. 13. 5. 7. 7. 7.Internodes (continued) Milled rice (continued) trade terms of. 30 type. 30 ciliation. 14 Multiple alleles. 7. 30 Opaque endosperm. 12. 22 shape. 14. 30 Milled rice. 15 axil. 28. 15. 7. 25. 21 seedling. 19. 20. 29. 11. 28. 22 38 . 10. 14. 14. 28 Mutant traits. 14 Pericarp. 5. 11. 30 Nerves. 14. 22 Pedicel. 5. 11. 7. 13 sperm. 7 Liguleless. 5. 28 pubescence. 30 Nucleus polar. 14 second. 19 weight. 13. 15. 25 extinction coefficient. 12 translucency. 22 weight. 28. 5. 14 Lower palea (See Lemma) Lutescent. 25 light transmission rate. 25 Ovary. 11. 7 elongated. 14. 28. 21. 13 Neckleaf. 7. 18 diameters. 24 Javanica. 7. 28. 28. 7. 10. 18 Microsporocytes. 14. 28. 17. 7. 7. 10 Nodal region (See Node) Nodal septum. 18 length. 13. 7. 19. 7. 7 length. 25 leaf area index. 30 Oryza glaberrima. 23. 21. 30 Macrosporocytes. 20 color. 5. 28 Neck (Also see Panicle base and collar). 20 form. 12 Pasting viscosity. 28 emergence. 5. 14 first (primary). 24. 22 Modifiers. 6. 23 Meiosis. 11. 28 number. 5. 10 Ligule. 20 Japonica. 14. 22 Milling. 31 Peduncle. 19 pubescence. 31 Panicle branches. 30 Lazy. 13. 30 exsertion. 14. 12. 13. 14. 14. 31 Panicle base. 27. 12. 5 Oryza sativa. 5. 7. 27. 7. 14. 27 Multiple pistils (See Polycaryoptic). 25 “Japonica’ subspecies. 13. 18. 18. 14. 28. 14 Parboiled rice. 5. 27 Leaf characters. 12 Midrib. 28 phenol staining. 24. 27. 20 insensitive varieties. 27. 14 Perianth. 12. 25. 30 Lodicules. 29. 22. 5. 14. 28 density. 12 Pale (See Palea) Palea. 15. 22 length. 7. 28 Maturity (Also see Growth duration). 5. 19 extra. 14. 27 Narrow blade. 14 Lamina (See Blade) Lax panicle. 11. 19 phenol staining.. 24. 7. 7.. 11. 14. 11. 30 color. 6. 20 Panicle axis. 31 color. 29 Lemma. 12 sheath. 13. 13. 14 senescence. 13. 28. 27. 12. 6. 6. 5 “second” (prophyll) . 30. 7. 11. 13. 25. 22 hardness. 12. 17. 14. 10 Palea inferior (See Lemma) Palea superior (See Palea) Palet (See Palea) Panicle. 27. 29 Mesocotyl. 19. 12. 10. 31 color. 15 Non-waxy. 31 Mucro. 17. 25 Leaf cushion (See Collar) Leaf spots. 14 Open hull. 25. 31 Photoperiod. 18. 13. 14. 19. 20 Internode elongation. 14 Ovary wall (See Pericarp) Ovule. 20 number. Node. 5. 18 shape. 7. 12. 15. 17.

18 Pulvinus panicle. 23 Threshing. 15. Pollen mother cells. 11. 31 Spikelet. 10. 14. 28. 6. 14 Tegmen. 15. 30. 22. 19. 11. 5 Roots (continued) seminal. 14 Protein content. 21 Pollen grains. 13. 11. 28. 29 Plumule. 21. 11. 11. 5 Shattering (shedding). 22. 31 Sheath (See Leaf sheath) Sheath joint (See Sheath pulvinus) Sheath pulvinus. 17. 12. 20 Physiologic diseases. 7. 11. 14. 19 auricles. 18 lemma and palea. 15. 28. 15. 24. 13 Starch. 25 Sterility. 12. 29 Stigma. 12. 30 Polyembryonic. 6. 29 Rachilln. 20 sensitive varieties. 13. 25. 28. 7. 11. 28. 15 Thermosensitive phase. 17. 27 Polycaryoptic. 31 Stamens. 15. 11. 13. 5 Subspecies. 18 Semidwarf. 28 fertility. 28. 15 primary (radicle) . test. 19 length. 15 Pollen fertility. 29 physiological diseases. 20. 12. 13 Striata. 13 Tiller. 13. 14. 13. 11. 24 Stem (See Culm) Sterile lemmas. 15 Seed coat (See Testa) Seedling. 25 ligule. 10. 19 Quantitative traits. 22 chemical properties of. 22. 18 blade surface. 15. 17. 15 Scales (See Lodicules) Scales for kernel dimensions and shapes. 23 physiological characters of. 5. 5 rootlets. 20 Resistance to chemicals. 18 blade margins. 10. 15. 17. 18 diseases. 14. 7. 23 Pigmentation. 18 cold water. 12 Pollen tube. 28 apiculus. 18. 22. 13. 27. 24 Spreading panicle branches. 30 shape. 14 . 5. 13 Rudimentary glumes (Also see Glumes). 5. 19 39 . 19 Rachis (See Panicle axis) Radicle. 23 insects. 20 sensitive phase. 5. 12. 12.Photoperiod (continued) optimum. 5 secondary. 11. 22 Scented kernel. 28. Polishing. 5 Seed (Also see Grain). 7. 15. 7. 13. 17. 11. 10. 11. 14 Pollination. 11. 30 Statistical techniques. 28 Red rice. 31 amylopectin. 25. 18 hairs. 6. 15 Section. 22. reaction to chemicals. 12. 13. 15 adventitious. 31 Pistil. 28 Style. 7. 15 color. 15 Sickles (See Auricles) Sinuous neck. 15. 31 clustering. 19. 19. 5. 14. 28 Series. 21. 11. 22 Proteinaceous matrix. 13. 11. 10. 19. 7. 23. 18. 27 Sheathing leaf (See Coleoptile) Shoot. 10. 27. 13. 14. 15. 13. 19 low air temperatures. 14. 22 Starch-iodine-blue. 28. 23 Physiologic races. 15 Rough rice (Also see Grain). 25.. 31 Scutellum. 7. 18 . 11. 20 number. 30 color. 11. 11. 31 Testa. 7. 19. 27. 20. 11. 14. 27 Reproductive phase. 11. 13. 15. 14. 10. 31 Prophyll (See Primary leaf) Prophyllum. 5. 5. 15 Proteins in leaves. 11. 23 Rhachis (See Panicle axis) Roiled leaf. 7. 31 Roots. 14 sheath. 30. 12. 7. 28 Recessive genes. 12. 13. 7. 25 Stool (See Tiller) Sub-family. 14. 12 Polyhusk (See Extra lemma) Potassium iodide-iodine solution. 23 diseases. 12 Polished rice (See Milled rice) . 19. 11. 21 Spodogram analysis. 25 Pubescence. 11. 27 color. 12. 6. 25. 15. 12. 15 Ratoon crop. 27. 29 Straw. 14 Polish. 22 Starchy endosperm (Also see Endosperm). 29 Pleiotropic effect. 23 lodging. 20. 30 Polygenes. 12. 5. 14. 17. 13 Ratooning ability. 15. 31 amylose. 14. 14.

31 Xantha.Tiller (continued) primary. 31 White core. 28. 23 thermosensitive. 11 White stripe. 24 Vegetative organs. 28. 20. of Agriculture. 7. 11. 31 Very hairy. 28. Yield components. 5. 31 U. hull. 28. 28. 31 Yield. 23 lowland. 31 White back. 22. 7. 11. 31 Yellow leaf. 24 . 22 White belly. 15 tertiary. 22 Upper palea (See Palea) Vagina (See Leaf sheath) Valve (See Lemma) Valvule (See Palea) Varietal catalog. 16 Tipburn yellow. 31 Trade terms. 13 40 . 28 Waxy endosperm. 31 Tribe. 23 Varieties (continued) saline-resistant. 25 Varieties. 28. 5-7 Verticillate rachis. 21. 28. 13 Twisted kernel. 31 Undulate rachis. 31 White hull. 23 photoperiod-sensitive. Department. 31 White streak. 24 Zebra stripe. 31 Zygote. 28. 23 floating. 20 upland. 5 Triploid. 16 secondary. 7. 28. 5 classification of. 30 Virescent. 11. 25 “deep-water”. 22. 15.S. 31 Viridis. 11. 12 Triangular. 23 Variety-groups.