Gill raker structure and selective predation on zooplankton by particulate feeding fish

The relationship between the gill raker structure of planktivorous fish (number, distance between gill rakers and length) and selective feeding on different species and size classes of zooplankton was studied. Gill raker structure was measured for brown trout Salmo trutta , Arctic charr Salvelinus alpinus , whitefish Coregonus lavaretus , roach Rutilus rutilus , bleak Alburnus alburnus , and three-spined stickleback Gasterosteus aculeatus . All species are facultative planktivorous fish and occur commonly in Scandinavian lakes. The effect of gill raker structure was studied by comparing prey found in fish stomachs with the availability of zooplankton from several lakes. Gill raker length and distance were significantly correlated with fish length. Although gill raker structure differed among species, all fish species selected the larger zooplankters. The minimum size of cladoceran species found in fish stomachs was much smaller than the distance between gill rakers. Despite great differences in gill raker spacing, the minimum size ingested of Daphnia galeata and Bosmina longispina was similar for all predators. The hypothesis that small zooplankton are strained and retained by the gill rakers in particulate feeding planktivorous fish, particularly in salmonids and roach, is rejected.     

Experimental research on the feeding of bream (Abramis brama L.) and white bream (Blicca bjoerkna L.)

Summary The feeding efficiency (the amount of food organisms (chironomids) eaten per unit time and weight raised to the power 0.8) of bream and white bream was determined in relation to the grain size of the substrate, in which the food was presented. Sand was used as substrate with a grain size up to 1500 , divided in eight fractions. The bream used had lengths of 14 and 33 cm and the white bream had a length of 21 cm.The experiments showed quite clearly that the grain size of the substrate influenced the feeding efficiency of all these fishes. An increasing grain size resulted in an decreasing feeding efficiency, because only up to a definite grain size the substrate can be filtered through the gill rakers. If the grain size is larger, the substrate cannot be sieved and is spit out. The size at which this switch occurred was different for all the fishes. For the small bream it was 350 , for the large bream 500 , and for the white bream 600 .Comparing the feeding efficiencies for a grain size which can be sieved by all these fishes, no significant difference was found between large bream and white bream. The small bream, however, was less efficient. Probably this difference is caused by the different sucking capacities of small and large fish. The small bream cannot in one time suck in the chironomids presented at 1 cm depth, while large bream and white bream can. So both grain size and depth at which the food was presented differentially determined the feeding efficiency of these fishes.     

Ecomorphological diet predictions: an assessment using inland silverside (Menidia beryllina) and longear sunfish (Lepomis megalotis) from Lake Texoma

The functions of an organisms morphological structures indicatethe organisms potential resource use (fundamental niche). Whilethis information often is also used to predict differences inactual resource use (realized niche) among individuals or species,such predictions may not be accurate because the maximum abilitiesmay not be useful to the organism under specific conditions or inspecific environments. We investigated the importance of sixpreviously studied morphologically based performanceabilities/constraints in structuring the diet of Menidiaberyllina (inland silverside) and Lepomis megalotis (longearsunfish) in Lake Texoma, a reservoir in the Red River basin(Oklahoma-Texas, USA). Of the six morphological characteristicsmeasured (number of gill rakers, length of gill rakers, spacebetween gill rakers, eye lens diameter, mouth size, mouthprotrusibility), only one characteristic for M. beryllina(mouth size) and three for L. megalotis (space betweenrakers, mouth size, and raker length) correlated with the gutcontents as predicted by previous functional morphology studies.This indicates that caution should be exercised when makinguntested predictions about the ecology of an organism based on itsfunctional morphology.     

Making a master filterer: Ontogeny of specialized filtering plates in silver carp (Hypophthalmichthys molitrix)

Filter feeding fishes possess several morphological adaptations necessary to capture and concentrate small particulate matter from the water column. Filter feeding teleosts typically employ elongated and tightly packed gill rakers with secondary bony or epithelial modifications that increase filtering efficiency. The gill rakers of Hypophthalmichthys molitrix, silver carp, are anatomically distinct from and more complex than the filtering apparatus of other teleostean fishes. The silver carp filtering apparatus is composed of biserial, fused filtering plates used to capture particles ranging in size from 4 to 80 μm. Early in ontogeny, at 15–25 mm standard length (SL), silver carp gill rakers are reminiscent of other more stereotypical teleostean rakers, characterized by individual lanceolate rakers that are tightly packed along the entirety of the branchial arches. At 30 mm SL, secondary epithelial projections and concomitant dermal ossification begin to stitch together individual gill rakers. During later juvenile stages, dermal bone further modifies the individual gill rakers and creates a bony scaffold that supports the now fully fused and porous epithelium. By adulthood, the stitching of bone and complete fusion of the overlying epithelium creates rigid filtering plates with morphologically distinct faces. The inner face of the plates is organized into a net‐like matrix while the outer face has a sponge‐like appearance comprised of differently sized pores. Here, we present morphological data from an ontogenetic series of the filtering apparatus within silver carp. These data inform hypotheses regarding both how these gill raker plates may have evolved from a more basal condition, as well as how this novel architecture allows this species to feed on exceedingly small phytoplankton, particles that represent a greater filtering challenge to the typical anatomy of the gill rakers of fishes.     

Genetic differentiation in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka

The genetic and environmental basis for polymorphism in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka, was investigated. Analysis of 30 full sib families involving pure types and reciprocal hybrids revealed that the variation was partitioned significantly among families within cross types and among cross types in both traits. As in the wild, kokanee displayed more gill rakers than sockeye; reciprocal hybrids displayed intermediate counts. Gill raker length also varied markedly among cross types, with pure sockeye displaying 19% longer gill rakers than comparable sized kokanee. This difference was in the opposite direction predicted, given the common positive association between gill raker number and length in sympatric morphs of the same species in fishes. Gill raker number and length were generally not correlated within cross types, suggesting independent divergence of the traits. The results are discussed in relation to genetic and trophic divergence of the morphs and to factors selecting for differentiation in the two gill raker traits.     

The role of predation and competition in determining the distribution of common bream,roach and white bream in Dutch eutrophic lakes

Synopsis Most of the lakes in The Netherlands are turbid and without vegetation. This is regarded as the result of increasing eutrophication within the last decades. Under these conditions common bream, roach, and white bream are the most common cyprinids. In six shallow (1–3 m), wind exposed lakes the abundance of common bream and roach was linked to the abundance of pikeperch; bream dominated when pikeperch was abundant, but when the latter species was rare, roach was dominant and the biomass of bream was reduced. The biomass of white bream was always relatively low. In lake Tjeukemeer the distribution of roach < 20 cm fork length (FL) and bream was also related to the distribution of pikeperch. Only roach > 20 cm FL managed to coexist with pikeperch in the open water area, whereas roach < 20 cm was confined to the littoral zone where pikeperch was nearly absent. Bream occurred mainly in the open water and avoided the littoral zone where it competed with roach. White bream occupied an intermediate position, occurring in relatively low density both in the littoral zone and in open water. The importance of predation and competition in determining the distribution and abundance of roach and common bream in the eutrophic lakes of The Netherlands is discussed.     

Diversification, Sympatric Speciation, and Trophic Polymorphism of Arctic Charr, Salvelinus Alpinus Complex, in Transbaikalia

In northern Transbaikalia, independently evolving landlocked populations of Arctic charr are found in mountain lakes. To assess the diversity of charr in this region, speciation modes involved in the evolution of charr forms, and the role of trophic polymorphism in their divergence, we studied the morphology and feeding of dwarf, small, and large forms of Arctic charr from a number of Transbaikalian lakes. Meristic data on charr from five lakes support the earlier conclusion that the three forms do not represent separate lineages but have independently diverged in sympatry in each of the lakes. In 10 lakes, the dwarf form showed varying degrees of differentiation from normal (small and large) charr in meristic characters (up to morphologically distinct and presumably reproductively isolated groupings), which is viewed as various levels of sympatric divergence. Indexes of gill raker length in fish from 20 lakes vary among populations of both dwarf and normal charr, with forms having short and long rakers being sympatric in some of these lakes. However, the index can be used only for comparing charr of different forms up to about 32cm fork length (FL) because it is strongly negatively correlated with size in larger fish. The study of charr diets in 21 lakes indicates that large charr are piscivorous whereas dwarf and small charr feed on a wide range of invertebrates, partitioning these resources in different ways. Planktivores, including very specialized ones, and non-planktivores (benthic feeders, insectivores) can be identified within the small and dwarf forms. The proportion of plankton in the diets of dwarf and small charr is positively correlated with the number and length of gill rakers while the proportion of benthos is negatively correlated. Allopatric planktivorous and non-planktivorous small charr differ in body proportions; parallel emergence of such morphotypes in different parts of the range is a characteristic feature of the Salvelinus alpinus complex.     

Feeding of benthophagous fish in the Chogray Reservoir

Data on feeding of the bream Abramis brama, roach Rutilus rutilus, and white bream Blicca bjoerkna in Chogray Reservoir in 2010 and 2012 are presented. The main diet of bream consists of Chironomidae, and the diet of roach and white bream consists of the bivalve mollusk Dreissena polymorpha.     

Experimental evidence for trait utility of gill raker number in adaptive radiation of a north temperate fish

North temperate fish in post‐glacial lakes are textbook examples for rapid parallel adaptive radiation into multiple trophic specialists within individual lakes. Speciation repeatedly proceeded along the benthic–limnetic habitat axis, and benthic–limnetic sister species diverge in the number of gill rakers. Yet, the utility of different numbers of gill rakers for consuming benthic vs. limnetic food has only very rarely been experimentally demonstrated. We bred and raised families of a benthic–limnetic species pair of whitefish under common garden conditions to test whether these species (i) show heritable differentiation in feeding efficiency on zooplankton, and (ii) whether variation in feeding efficiency is predicted by variation in gill raker numbers. We used zooplankton of three different size classes to investigate prey size dependency of divergence in feeding efficiency and to investigate the effect strength of variation in the number of gill rakers. Our results show strong interspecific differences in feeding efficiency. These differences are largest when fish were tested with the smallest zooplankton. Importantly, feeding efficiency is significantly positively correlated with the number of gill rakers when using small zooplankton, also when species identity is statistically controlled for. Our results support the hypothesis that a larger number of gill rakers are of adaptive significance for feeding on zooplankton and provide one of the first experimental demonstrations of trait utility of gill raker number when fish feed on zooplankton. These results are consistent with the suggested importance of divergent selection driven feeding adaptation during adaptive radiation of fish in post‐glacial lakes.     

Temporal variation in feeding morphology and size-structured population dynamics in fishes

Considerable variation in morphology associated with resource use is a classic example of local adaptation to the environment. We demonstrate that a temporal change in feeding morphology might occur within a population. In a 5-year natural field experiment, we estimated gill raker morphology, resource density and population dynamics of the roach and its competitor, the perch. Despite a variation in density of zooplankton resources and perch across years, no change in roach density was observed. However, gill raker morphology in roach covaried with size structure of the zooplankton resource across years. A laboratory experiment confirmed that gill raker morphology has a great effect on roach foraging efficiency on zooplankton and that there is a functional trade-off with regard to zooplankton foraging. We suggest that the response in gill raker structure is an adaptation to deal with the rapid population dynamics of zooplankton, which are in turn mediated by changes in the size structure of the competing perch.     

Ecological consequences of food partitioning for the fish population structure in a eutrophic lake

In the highly eutrophic lake, Frederiksborg Slotssø, the diet composition of the bream (Abramis brama L.) and roach (Rutilus rutilus L.) populations was examined during three periods with different food availability. The length range of bream and roach was 9–34 cm (TL) and 5–18 cm (TL), respectively. The relative food composition was examined for 2 cm and 1 cm length intervals of bream and roach, respectively. During all three periods, bream shifted from benthic cladocerans (Alona sp.) to zooplankton and chironomids within a transitional length of 15.0–20.0 cm. These foodshifts were coupled with a change in feeding behaviour from particulate to filter feeding. The biomass of chironomids was too low to sustain the consumption of larger bream (>20.0 cm) which initiated feeding in the pelagic zone even in periods when the mean length and biomass of the preferred zooplankton, Daphnia cucullata, were low. In contrast to bream, roach fed mainly on zooplankton. With increasing size, roach progressively shifted to larger zooplankton species due to the increasing mesh size of their branchial system. The importance of benthic animals in the diet of roach was minor due to low feeding efficiency on prey buried in the sediment. Detritus appeared in the diet of bream and roach in periods of low availability of animal food items. Feeding on detritus may provide an energetic advantage to bream and roach and increase the carrying capacity for these species in lakes, where detritus is highly abundant. Especially for the larger fish due to the decrease in their relative metabolic demands. However, the ability of bream to filter feed and with increasing size to retain food items smaller than those retained by roach may be the main mechanism for the dominance of bream over roach in highly eutrophic lakes.     

New insights into the migration and habitat use by bream and white bream in the floodplain of the River Rhine

Bream Abramis brama , white bream Blicca bjoerkna and roach Rutilus rutilus all undertook spawning migrations from the River Rhine into oxbows. Adult white bream and roach returned to the river after spawning. This indicates an annual change of habitat allowing the use of optimal conditions in oxbows for spawning and growth of juvenile fish, and the use of feeding grounds rich in benthos in the main river by the adults. In contrast, adult bream were still present in large numbers in the oxbows over the summer. Juvenile bream (0+ and 1+) migrated to the main river. Bream aged 2+ to 4+ were not found in the oxbows, whereas they occurred frequently in the river. Thus bream show ontogenetic changes in their use of the main river and oxbows. The young fish spend their long juvenile phase in the river after leaving the oxbows. Reproduction in oxbows is ensured by mature fish migrating in for a short period from the river as well as by other adults (≥5+) that live permanently in the oxbows. The strong tendency of adult bream to develop permanent stocks in the oxbows gives this species a reproductive advantage, since many oxbows are accessible to Rhine fish only occasionally in the present hydrological situation.     

Integration of pecking,filter feeding and drinking mechanisms in waterfowl

This paper is one of several contributions in a series, illustrating the application of a specific deductive methodology to explain diversity of form. The methodology facilitates the explanation of feeding morphologies in various ducks as a transformation of the mallard's feeding design maximized for specific proportions of performance that are contributed by pecking and filter feeding mechanisms.The earlier described anatomy and formal analyses of the three mechanisms in the mallard served as the initial conditions used in simulation models. Four elements of the feeding system were chosen that play a major role in all three mechanisms. For each element, the main parameter was selected: storage capacity of the rostral mouth cavity, transport capacity of the rostral mouth tube, storage capacity of the caudal mouth cavity and transport capacity of the caudal mouth tube. The boundary conditions for the simulation were determined from internal organismic constraints. The total food uptake of the mallard was regarded as the function to be maximized. This object function is the summation of the food uptake by one second of pecking and one second of filter feeding. The drinking mechanism was shown not to interfere, since that mechanism operates sufficiently whenever the pumping mechanism works properly.The object function, made up by the pecking and filter feeding performances was graphed. From these graphs a morphospace was developed: the region within which modifications of the mouth design are feasible. This procedure allowed examination of the general hypothesis that different modifications of one design for a complex multi-role system are explainable from differences in proportions of the functional performance contributed by each of the roles.Two predictions were evaluated more specifically: 1) If filter feeding performance must increase for a specific change in total food uptake, the volume of the rostral mouth cavity must increase; this requires widening and lengthening of the rostral maxillar portion and also a phase shift in jaw and lingual motion patterns, increasing the stroke volume. 2) If pecking performance must increase, the transport capacity of the rostral mouth tube must increase; this requires shortening of the maxillar mid portion. These two predictions regarding change in mouth morphology were borne out by shovelers and tufted ducks, respectively.The deductive analysis of the feeding system's morphospace allowed the explanation of the wide diversity of forms in the duck mouth designs as emerging from the great radiation capacity of the ancestral duck's mouth design: it is a versatile, potentially multi-role system in which pecking, drinking and filter feeding mechanisms are mutually very tolerant and, by epiphenomena, includes the elements of a grazing mechanism. In addition, the deduced morphospace was shown to explain phenotypic plasticity, since it explains the occurrence of two morphotypes that develop due to habitat discrimination: one type that has been forced to filter feed, and one type that feeds by pecking.Address reprint requests to Dr. G.A. Zweers     

Variation in number of vertebrae and gill rakers of sockeye salmon,Oncorhynchus nerka,in North America

Synopsis The latitudinal, regional, and annual variation in number of vertebrae and number of gill rakers present in sockeye salmon, Oncorhynchus nerka, stocks in North America was examined. Stocks in more northern areas had higher numbers of vertebrae and gill rakers than did those in more southern ones. Significant annual variability in the frequencies of these meristic characters within stocks was observed. When stocks were grouped into three regions (southern, central, and northern), heterogeneity in vertebral and gill raker frequencies was greater among regions than among stocks within the regions. Similarly, heterogeneity was greater among stocks than among sampling years within stocks. Differences in vertebral and gill raker frequencies are only useful for stock identification of sockeye salmon on a broad regional basis.     

Coexistence and origin of trophic ecotypes of pygmy whitefish,Prosopium coulterii,in a south‐western Alaskan lake

Ecologically, morphologically and genetically distinct populations within single taxa often coexist in postglacial lakes and have provided important model systems with which to investigate ecological and evolutionary processes such as niche partitioning and ecological speciation. Within the Salmonidae, these species complexes have been well studied, particularly within the Coregonus clupeaformis–C. laveratus (lake and European whitefish, respectively) group, but the phenomenon has been less well documented in the other whitefish genera, Prosopium and Stenodus. Here, we examined the morphology, feeding biology and genetic structure of three putative forms of the pygmy whitefish, Prosopium coulterii (Eigenmann & Eigenmann, 1892), first reported from Chignik Lake, south‐western Alaska, over 40 years ago. Field collections and morphological analyses resolved a shallow water (< 5 m depth) low gill raker count form (< 15 first arch gill rakers), a deepwater (> 30 m), low gill raker form and a deepwater, high gill raker count (> 15 gill rakers) form. The two low gill raker count forms fed almost exclusively on benthic invertebrates (mostly chironomids), while the deepwater, high gill raker count form fed almost exclusively on zooplankton; differences in diet were also reflected in differences both in δ¹³C and δ¹⁵N stable isotopes. All three forms were characterized by the same major mitochondrial DNA clade that has been associated with persistence in, and postglacial dispersal from, a Beringian glacial refugium. Analysis of variation at nine microsatellite DNA loci indicated low, but significant differentiation among forms, especially between the two low gill raker count forms and the high gill raker count form. The extent of differentiation along phenotypic (considerable) and genetic (subtle) axes among the Chignik Lake forms is similar to that found among distinct taxa of Prosopium found in pre‐glacial Bear Lake (Utah–Idaho, USA) which is probably at least ten times older than Chignik Lake. Our analyses illustrate the potential for the postglacial differentiation in traits subject to divergent natural selection across variable environments.     

Trophic morphology features allow Astyanax endemic species coexistence in a Neotropical river system

Resource partitioning has an essential role in interspecific relations, especially in congener species, which share many morphological traits. In some places, small characids coexist through resource partitioning, which may reduce their interspecific competition. Astyanax species (e.g., Astyanax minor, Astyanax gymnodontus and Astyanax bifasciatus), for example, coexist in different water bodies from the Iguaçu River basin. These species have high phenotypic plasticity and many morphologic specializations that allow them to live in different habitats. Based on evidences that these species modified their feeding habits because of changes in resource availability in Iguaçu River, this study tested two hypotheses: (a) there are differences in head morphology, number of teeth and number of gill rakers among the species of Astyanax; and (b) there are differences in gill arch and gill raker morphology among the species of Astyanax, which may favour their trophic resource partitioning in Iguaçu River. The head morphological traits and quantitative morphological characters were summarized in a principal coordinates analysis (PCoA), and the analysis of similarities (ANOSIM) showed significant differences among species. Gill morphological measurements were analysed through analysis of covariance (ANCOVA), and it also showed significant differences in gill arch and gill raker morphology among species. Therefore, the analysis of ecomorphological traits related to trophic habits revealed some differences that may suggest a tendency of reducing competition for trophic resources in the Iguaçu River basin.     

Effects of root herbivory by an insect on a foliar-feeding species,mediated through changes in the host plant

Summary The effects of root herbivory by larvae of the scarabaeid, Phyllopertha horticola, on the growth of Capsella bursa-pastoris were examined. Individuals of Aphis fabae were reared on the leaves to determine what effect, if any, root feeding has on the performance of this insect. The experiment was conducted under two watering regimes (low and high). Low watering and root feeding caused water stress in the plants and this was reflected in a reduction in vegetative biomass and an increase in the proportion of material allocated to reproduction. Supplying plants with ample water in the high treatment enabled the water stress caused by root herbivory to be offset, but not completely overcome. Low watering and root feeding caused an increase in aphid weight and growth rate, while root feeding also increased fecundity and adult longevity. These effects are attributed to an improvement in food quality, measured by total soluble nitrogen, and caused by amino acid mobilization due to the water stress. The implications of these results in agricultural and ecological situations are discussed.     

Trophic polymorphism, habitat and diet segregation in Percichthys trucha (Pisces: Percichthyidae) in the Andes

Divergent natural selection affecting specific trait combinations that lead to greater efficiency in resource exploitation is believed to be a major mechanism leading to trophic polymorphism and adaptive radiation. We present evidence of trophic polymorphism involving two benthic morphs within Percichthys trucha , a fish endemic to temperate South America. In a series of lakes located in the southern Andes, we found two morphs of P. trucha that could be distinguished on the basis of gill raker length and five other morphological measures, most of which are likely associated with the use of food resources. The differences were consistent across all lakes examined, and were correlated with habitat use and diet. Individuals with longer gill rakers were more abundant in the littoral zone (littoral morph) while the short gill-raker morph was more abundant at 10 m depth and deeper (deep benthic morph). Both morphs fed primarily on benthic invertebrates, but the littoral morph fed more on larval Anisoptera than did the deep benthic morph. Phenotypic correlations among traits were high for the littoral morph, but low and non-significant for the deep-benthic morph. We suggest that gill raker length may influence the relative efficiency of suction feeding for the two morphs. This is the first evidence of trophic polymorphism in fishes from temperate South America.     

The role of gill raker number variability in adaptive radiation of coregonid fish

Gill raker divergence is a general pattern in adaptive radiations of postglacial fish, but few studies have addressed the adaptive significance of this morphological trait in foraging and eco-evolutionary interactions among predator and prey. Here, a set of subarctic lakes along a diversifying gradient of coregonids was used as the natural setting to explore correlations between gill raker numbers and planktivory as well as the impact of coregonid radiation on zooplankton communities. Results from 19 populations covering most of the total gill raker number gradient of the genus Coregonus, confirm that the number of gill rakers has a central role in determining the foraging ability towards zooplankton prey. Both at the individual and population levels, gill raker number was correlated with pelagic niche use and the size of utilized zooplankton prey. Furthermore, the average body size and the abundance and diversity of the zooplankton community decreased with the increasing diversity of coregonids. We argue that zooplankton feeding leads to an eco-evolutionary feedback loop that may further shape the gill raker morphology since natural selection intensifies under resource competition for depleted prey communities. Eco-evolutionary interactions may thus have a central role creating and maintaining the divergence of coregonid morphs in postglacial lakes.     

Spontaneous chlorophyll mutants of Pennisetum americanum: Genetics and chlorophyll quantities

Summary Thirteen spontaneously occurring chlorophyll deficient phenotypes have been described and their genetic basis was established. Ten of these — white, white tipped green, patchy white, white virescent, white striping 1, white striping 2, white striping 4, fine striping, chlorina and yellow virescent showed monogenic recessive inheritance and the remaining three — yellow striping, yellow green and light green seedling phenotypes showed digenic recessive inheritance. The genes for (i) white tipped green (wr) and yellow virescent (yv) and (ii) patchy white (pw) and white striping 1 (wst 1) showed independent assortment. Further, the genes for white (w), white tipped green (wr) and yellow virescent (yv) were inherited independently of the gene for hairy leaf margin (Hm).In the mutants — white tipped green, patchy white, white striping 1, white striping 2, fine striping, chlorina, yellow virescent, yellow striping, yellow green and light green phenotypes total quantity of chlorophyll was significantly less than that in the corresponding controls, while in white virescent there was no reduction in the mature stage. For nine of the mutants the quantity of chlorophyll was also estimated in F₁'s (mutant x control green). In F₁'s of six of the mutants — white tip, patchy white, chlorina, yellow virescent, fine striping and yellow striping the quantity of chlorophyll was almost equal to the wild type. In the F₁'s of three of the mutants — white striping 1, white striping 2 and light green an intermediate value between the mutant and wild types was observed. In yellow virescent retarded synthesis of chlorophyll, particularly chlorophyll a was observed in the juvenile stage. Reduced quantity of chlorophyll was associated with defective chloroplasts. In the mutants — white tipped green, white virescent, fine striping, chlorina, yellow striping, yellow green and light green defective plastids were also observed. In patchy white secondary destruction of chlorophylls and the presence of defective plastids were found to be associated with reduced chlorophyll quantity at maturity.Paper chromatographic studies of leaf flavonoids revealed some variation between the inbreds, but there were three common spots, 7, 8 and 9, except for PDP in which the spot 8 was absent. Chlorophyll deficient mutants differed from their respective controls in the absence of one or more of the spots present in the controls and in the presence of new spots in some of the mutants.Most of the chlorophyll mutants showed higher survival rate in the Kharif season than in Rabi season which was attributed to the higher mean day temperature and longer day light period in the Kharif season than in Rabi season.