Phenotypic and Genetic Correlations Among Floral Traits in Two Species of Thalictrum

Save this PDF as:
 WORD  PNG  TXT  JPG

Size: px
Start display at page:

Download "Phenotypic and Genetic Correlations Among Floral Traits in Two Species of Thalictrum"

Transcription

1 gen, non-protein nitrogen and nucleic acids during wheat grain development. Aust J Physiol 6: Evers AD, Development of the endosperm of wheat. Ann Bot 34: Kidd AD, Francis D, and Bennett MD Replicon size, mean rate of DNA replication and duration of cell cycle and its component phase in eight monocotyledonous species of contrasting DNA values. Ann Bot 59: Kowles RV and Phillips RL, DNA amplification patterns in maize endosperm nuclei during kernel development. Proc Natl Acad Sci USA 82: Kowles RV, Srienc F, and Phillips RL, Endoreduplication of nuclear DNA in the developing maize endosperm. Dev Genet 11: Nagl W, Polytene chromosomes of plants. Int Rev Cytol 73: Phillips RL, Wang AS, and Kowles RV, Molecular and developmental cytogenetics of gene multiplicity in maize. Stadler Genet Symp 15: Ramachandran C and Raghavan V, Changes in nuclear DNA content of endosperm cells during grain development in rice (Oriza sativa). Ann Bot 64: Corresponding Editor: Prem P. Jauhar Phenotypic and Genetic Correlations Among Floral Traits in Two Species of Thalictrum S. L. Davis The evolution of dioecy in plants is expected to be followed by sex-specific selection, leading to sexual dimorphism. The extent of the response to selection depends on the genetic covariance structure between traits both within and between the sexes. Here I describe an investigation to determine phenotypic and genetic correlations between reproductive traits within cryptically dioecious Thalictrum pubescens and within morphologically dioecious T. dioicum. Females of T. pubescens produce flowers having stamens and pistils, appearing hermaphroditic. Genetic correlations were estimated as family-mean correlations among paternal half-sib families. Positive phenotypic and genetic correlations between parts of the same reproductive organs, as the anther and filament of the stamen, indicate developmental associations between these traits in both species. Negative genetic correlations were detected between pistil number and size of reproductive organs in T. dioicum and showed the same direction, but not significance, in T. pubescens. There was a negative phenotypic correlation between the number of stamens and the number of pistils within female flowers of T. pubescens. Within T. pubescens, there was a positive genetic correlation between the number of stamens in males and the number of pistils in females, indicating that floral evolution in males and females may not be independent in this species. Avoidance of inbreeding depression and resource reallocation have long been considered primary factors driving the evolution of gender dimorphism and/or dioecy from cosexuality in plants, yet their relative contributions to this pathway are still argued ( Bawa 1980; Charlesworth and Charlesworth 1978; Charnov et al. 1976; Darwin 1877; Willson 1979). Whatever the nature of selection driving the evolution of separate sexes, the response to that selection will depend on the relationship, particularly the genetic correlations, between male and female reproductive traits ( Lande 1980). Theoretical predictions as to what the genetic relationship between reproductive traits might be and what empirical studies have shown have not always coincided. For example, a major assumption underlying theoretical models of sex allocation is the existence of a fixed amount of resources available for reproduction that can be allocated in various ways among the different sexual functions (Charlesworth and Morgan 1991). This leads to the expectation of negative genetic correlations between male and female traits within cosexuals. Such negative correlations between male and female traits should help augment selection toward sexual specialization among individuals, as individuals that allocate more to male function conversely allocate less to female function (Mazer et al. 1999). Yet studies have often found positive correlations in allocation to male and female functions (Ashman 1999; Campbell 1997; Elle 1998; Fenster and Carr 1997; O Neil and Schmitt 1993). Positive correlations between traits can be caused by greater genetic variation among individuals in resource acquisition than in allocation (Houle 1991), so individuals with more resources available would allocate more to both male and female traits. Selection can also shape genetic correlations between two traits that interact together to perform a given function. If functional integration between traits increases an individual s fitness, selection would act to increase the positive correlation between those traits (Conner 1997; Conner and Via 1993). Once dioecy has been established, further changes between the sexes in resource allocation patterns are expected to occur because of sex-specific selection (either sexual or natural selection (Geber 1995), leading to sexual dimorphism (Cox 1981; Lloyd and Webb 1977; Meagher 1994; Willson 1979). However, positive genetic correlations between traits in each sex would limit the independent evolution of those traits in males and females (Lande 1980; but see Cheverud et al. 1985). For example, selection against a particular trait may be countered by selection favoring that same trait in the opposite sex. Positive genetic correlations between the sexes are predicted because when dioecy first arises, sexual dimorphism should be low and homologous characters in the two sexes should be under the control of the same or overlapping sets of genes in each sex (Meagher 1992, 1994). One possible example of this constraint on the evolution of sexual dimorphism may be found in dioecious species in which at least one of the sexes produces full size, yet nonfunctional, sexual organs of the opposite sex, making them appear hermaphroditic. This condition is known as cryptic dioecy (Mayer and Charlesworth 1991). The maintenance of cryptic dioecy could be explained by the existence of positive genetic correlations between the sexes so that, for example, selection for resource reallocation and reduction of anthers in females would be countered by selection favoring anthers in males. Hence the study of the genetic relationship between reproductive traits in cryptically dioecious species and their relatives offer a unique opportunity to better understand factors influencing breeding system evolution and sexual dimorphism. In this article I describe an investigation to determine the relationship between reproductive traits within the cryptically dioecious species Thalictrum pubescens and its dioecious relative T. dioicum. As with most cryptically dioecious species, T. pubescens is morphologically androdioecious but functionally dioecious. Females produce flowers that have both stamens and pistils, making them appear hermaphroditic. T. dioicum is functionally and morphologically dioecious. Females of T. pubescens appear to invest the same amount of resources per stamen as males. Furthermore, a possible trade-off between stamen and pistil production within female flowers indicates that females may be maintaining stamen production at the expense of pistil production, and hence potential seed production ( Davis SL, in preparation). Therefore there should be strong selection against producing stamens in females. Davis (1997) found that females of T. pubescens whose stamens Downloaded from at Pennsylvania State University on March 6, 2014 Brief Communications 361

2 had been removed did not suffer reduced seed set due to a lack of adequate pollination by insects, most likely because this plant is also pollinated by wind. Hence stamens in females of this species are not likely maintained as a reward for pollinators. Consequently I have investigated the genetic relationship among the reproductive traits in this species to help understand why females are morphologically hermaphroditic. Materials and Methods Study Species T. pubescens and T. dioicum are members of the Ranunculaceae. Sex determination in Thalictrum sp. appears to be under nuclear control, with males serving as the heterogametic sex (reviewed in Meagher 1988). Both species lack petals and nectaries ( Boivin 1944). Sepals are small and whitish and fall off quickly as the flower ages. T. pubescens is a summer flowering perennial that grows in rich woods, low thickets, swamps, wet meadows, and stream banks. Females of T. pubescens are functionally unisexual, but morphologically hermaphroditic: females produce pistillate flowers with an average of 12 pistils with uniovulate ovaries and 8 stamens containing inaperturate pollen. Males produce only staminate flowers with an average of 38 stamens and no vestigial pistils (Davis SL, in preparation). Both wind and insects have been reported to pollinate this species (Davis 1997; Kaplan and Mulcahy 1971; Keener 1976). T. dioicum is an early spring flowering perennial that grows in rich rocky woods, ravines, and alluvial terraces ( Keener 1976). Flowers of both sexes are both functionally and morphologically unisexual. This species is solely wind-pollinated and shows characteristics associated with this mode of pollination: stamens of males are pendulous and pistils of females are upright. Field sites for the populations to be used in the study are Cedar Bluffs State Nature Preserve located near Bloomington, Indiana, for T. dioicum and a small unused field located near Fort Hill State Historical Park near Hillsborough, Ohio, for T. pubescens. Phenotypic and Genetic Correlations When character traits to be correlated cannot be measured on the same individual (such as individuals growing in different environments), genetic correlations cannot be estimated directly ( Via 1984). Genetic correlations between the sexes are directly analogous to genetic correlations between environments in that the two sexes represent different physiological environments (Meagher 1992). An approximation to the genetic correlation is the correlation of family means. The estimation of covariation of family means (cov m ) contains both the covariance among families and a fraction of the within-family error variance, so cov m cov among (1/n)cov within, where n is the number of individuals per family. Hence the standard family mean correlation is expected to estimate the true value as the number of individuals per family increases (Via 1984). It is safe to assume that any significant negative correlations within or among taxa represent real genetic constraints, because any bias due to environmental effects in the greenhouse would likely skew values in a positive direction (Fry 1993; Houle 1991; Mazer and Hultgård 1993). Seeds from 30 open-pollinated maternal plants of each species were collected from the field sites described above. Seeds of T. dioicum were collected in April 1994 and seeds of T. pubescens were collected in August Seeds of both species were planted in late September 1994 in trays of MetroMix soil and placed in a 4 C cold room for 6 weeks. Germination rate is low ( 30%), so large numbers of seeds by the handful per maternal plant were sown. Seeds from 28 maternal lines of T. dioicum and 21 maternal lines from T. pubescens germinated. Trays were then transferred to a mist room in a greenhouse at Indiana University. After the seeds germinated and seedlings reached a reasonable size (after 7 8 weeks), 2 15 seedlings per maternal line, depending on how many had germinated, were transplanted into 4 in. clay pots filled with a 3:1 mixture of soil and peat. Plants were placed in a greenhouse at the Botanical Experimental Field Station at Indiana University. After another 8 10 weeks of growth, the plants were transplanted into 8 in. pots, filled with a 3:1 mixture of soil and peat. When these plants flowered they were used to create paternal half-sib families in both species as described below. For T. dioicum, plants flowered in January March A total of 26 males, each from a different maternal line, were each crossed with three to five females from maternal lines different from each other and from the male being crossed. Seeds from each cross were collected and coldtreated, and germinated as described above. A maximum of 15 seedlings from each father mother combination was transplanted into 4 in. pots. Plants began to flower in February 1996, before they were transplanted, so they were kept in the 4 in. pots. On the first or second day of flowering, two flowers per plant were collected in 70% ethyl alcohol (ETOH) for measurements to be taken in the laboratory. Two flowers per plant seems an adequate number, as the within-plant variation among floral traits is much lower than the among-plant variation in floral traits. There were a total of 26 paternal families with an average of 12.4 individuals per family. Flowering rate was low in T. pubescens. Less than 20% of T. pubescens plants flowered in 1995, and another round of flowering started in August Although not ideal, by using plants in both rounds of flowering, I was able to use 17 males to pollinate two to three females each. Seeds from each paternal half-sib family were planted and cold-treated in September 1997 as described above. A maximum of 15 seedlings from each father mother combination of T. pubescens were transplanted into 4 in. pots in October To promote greater flowering rates, these plants were exposed to another round of cold-stratification by leaving the greenhouse unheated unless the temperature dropped below 40 F starting in early January In mid-march 1998, as the temperature increased, the plants were transplanted into 8 in. pots and moved into a greenhouse with the minimum temperature control turned off. Plants started flowering in early May. On the first or second day of flowering for each plant, two flowers were collected in 70% ETOH for measurements to be taken in the laboratory. One family failed to have any seeds germinate and one other failed to have any plants flower, giving a total of 15 families with an average of 16.2 individuals per family. Measurements on all flowers were taken using a micrometer on a dissecting microscope under 8 magnification. Traits measured were number of stamens, number of pistils, stigma length, ovary length, anther length, filament length, and sepal length. Phenotypic correlations were calculated by using the entire dataset for each species. Pearson product-moment correlation coefficients were calculated for each pair of variables within each sex. Because of the small number of plants that flowered at one time precluded a complete crossing design, the power of this experiment was limited. Variance components for maternal and paternal ef- 362 The Journal of Heredity 2001:92(4)

3 Table 1. Phenotypic and genetic correlations between floral traits in female T. dioicum Ovary length Number of pistils Sepal length Stigma length x 1 SD Ovary length (185) ** (184) ** (187) Number of pistils * (26) (182) (185) Sepal length (26) (26) ** (184) Stigma length (26) (26) (26) Phenotypic correlations are shown above the diagonal and family-mean correlations are shown below the diagonal. N is indicated in parentheses. Those correlations that are significant after correcting for multiple tests by using a sequential Bonferroni technique are shown in bold. fects could not be estimated and heritabilities for individual traits were not calculated. For estimates of genetic correlations, the mean of each trait was calculated for each paternal half-sib family. Pearson product-moment correlation coefficients were then calculated for each pair of traits among half-sib families. The number of data points could have been increased by collapsing the design and using full-sib families. However, these data points would not be independent of one another, since in the breeding design each mother was crossed with multiple fathers. The correlation between the number of stamens in males and the number of pistils in females in both species was calculated, as, in a sense, these characters represent homologous structures in the two sexes. In addition, for T. pubescens, correlations were calculated between the pairs of traits shared by males and females: number of stamens, stamen length, anther length. Because the correlations were calculated among paternal sibships, variation due to maternal effects should be minimal. Log-transformation of nonnormally distributed variables did not change the substance of the results, so the analyses on untransformed data are presented for ease of interpretation. For each group of correlations within each sex (i.e., within-female phenotypic correlations, within-male genetic correlations), significance levels were corrected for multiple tests using a sequential Bonferroni procedure (Rice Table ). The table-wise error rate was set at 0.05, which was then divided by the number of tests within the table. If the lowest P value within the table was less than this new adjusted value, that correlation was considered significant. The next lowest P value was then compared to a new value of 0.05 divided by the number of tests minus 1, and so on, until a nonsignificant test was reached. Results Phenotypic and Genetic Correlations between floral traits in male T. dioicum Number of stamen Anther length Filament length Sepal length x 1 SD Number of stamen (122) (122) (122) Anther length (26) ** (123) ** (123) Filament length (26) (26) ** (123) Sepal length (26) (26) (26) Phenotypic correlations are shown above the diagonal and family-mean correlations are shown below the diagonal. N is indicated in parentheses. Those correlations that are significant after correcting for multiple tests by using a sequential Bonferroni technique are shown in bold. Figure 1. Correlations among paternal family means between the number of stamens per flower produced in males and the number of pistils produced per flower in females: (A) Thalictrum dioicum; (B) Thalictrum pubescens. Thalictrum dioicum Two classes of phenotypic correlations were significant in both sexes. First, structures within organs were positively correlated: within the pistil, stigma length was positively correlated with ovary length (Table 1), and within the stamen, anther length was strongly correlated with filament length ( Table 2). Second, positive phenotypic correlations exist in both females and males between the size of sexual organs and the size of the sepals (Tables 1 and 2). No significant phenotypic correlations were associated with either the number of pistils or the number of stamens. There were no significant genetic correlations within male floral traits ( Table 2). A significant negative genetic correlation was present between ovary size and pistil number within female flowers ( Table 1). As ovary size increases, pistil number decreases. The genetic correlation between stamen number in males and pistil number in females was not significant (r 0.061, N 26, P.767; Figure 1A). Thalictrum pubescens Patterns of phenotypic correlations in T. pubescens were similar to those found in T. dioicum (Tables 3 and 4). Within the pistils of females, stigma length was positively correlated with ovary length ( Table 3), and within the stamens of both males and females, anther length was strongly correlated with filament length (Tables 3 and 4). Sepal length was positively correlated with the size of stamens and pistils in females and with stamen size in males. The number of pistils in females was negatively correlated with the size of the ovary and with the size of the stamen parts: flowers with large numbers of pistils had smaller stamens and pistils. In addition, there was a negative correlation between the number of stamens and the number of pistils within female flowers ( Table 3). Most of the genotypic correlations within females were in the same direction as the phenotypic correlations. However, after correcting for multiple tests, none of these correlations were significant ( Table 3). Within male flowers, anther length was again significantly correlated with filament Downloaded from at Pennsylvania State University on March 6, 2014 Brief Communications 363

4 Table 3. Correlations between floral traits in female T. pubescens Number of stamen Anther length Filament length Ovary length Number of pistils Sepal length Stigma length x 1 SD Number of stamen 0.241* (145) 0.386** (145) (147) 0.281** (147) 0.406** (148) (147) Anther length (15) 0.436** (145) 0.279** (144) 0.380** (144) 0.603** (145) 0.364** (144) Filament length (15) (15) 0.285** (144) 0.335** (144) 0.442** (145) 0.300** (144) Ovary length (15) (15) (15) 0.259* (147) 0.266** (147) 0.321** (147) Number of pistils (15) (15) (15) (15) (147) (147) Sepal length (15) (15) (15) (15) (15) 0.364** (147) Stigma length (15) (15) (15) (15) (15) (15) Phenotypic correlations are shown above the diagonal and family-mean correlations are shown below the diagonal. N is indicated in parentheses. Correlations that are significant after correcting for multiple tests by using a sequential Bonferroni technique are shown in bold. length, and again there was a trend for these traits to be positively correlated with sepal size (Table 4). In T. pubescens, there were no significant correlations between stamen characters in males and stamen characters in females as were expected. However, there was a positive correlation among families between the number of stamens and the number of pistils: sibships with females that have many pistils also have males that have many stamens (Figure 1B). Discussion The repeated detection of correlations between specific traits among closely related species can provide insights into selection patterns and the joint evolution of these traits (Mazer and Hultgård 1993). Despite the lack of power in the breeding design, the phenotypic and genetic correlations found in this study were similar in both species. These patterns of covariation are discussed below in terms of how they may contribute to evidence of genetic constraints on the independent evolution of floral traits within the genus. Correlations Within Flowers Positive genetic correlations between parts of the same structure, such as those found here between anther and filament length in the stamen and ovary and stigma Table 4. Phenotypic and genetic correlations between floral traits in male T. pubescens Number of stamen Anther length Filament length Sepal length x 1 SD Number of stamen (188) (188) (187) Anther length (15) ** (188) ** (187) Filament length (15) * (15) ** (187) Sepal length (15) (15) (15) Phenotypic correlations are shown above the diagonal and family-mean correlations are shown below the diagonal. N is indicated in parentheses. Those correlations that are significant after correcting for multiple tests by using a sequential Bonferroni technique are shown in bold. length in the pistil, are not surprising and most likely reflect strong developmental associations between these traits. These correlations might impose constraints on selection responses if, for example, males are selected to have longer filaments in order to aid in pollen dispersal in a windpollinated plant, but are under conflicting selection pressure to maintain a certain anther size. A second prevalent positive correlation was that of the phenotypic correlation between the size of the sexual organs and that of the sepals in both sexes of both species. As both species are wind pollinated and sepals fall off shortly after the flower matures, the perianth (in this case just sepals) most likely does not function in pollinator attraction, and hence does not play a factor in determining their size in males versus females. Two other possible hypotheses, however, might lead to the observed correlations. First, developmental associations between the corolla and stamens (androecium) may cause a positive correlation between these structures. Second, the role of the perianth in enclosing the reproductive structures in the bud may result in a correlation between the size of enclosed structures and the sepals or petals (Delph et al. 1996). Sepal size was phenotypically correlated with stamen size in both males and females of T. pubescens, but showed no correlation with pistil size in females, so it appears that in T. pubescens the data follows the prediction that developmental associations with stamens contribute to the determination of sepal size. In the morphologically dioecious T. dioicum, however, sepal size was phenotypically correlated with stamen size in males and pistil size in females, so a developmental relationship with anthers alone cannot account for sepal size and the protective function of the sepals may be playing a more prominent role. There were no significant negative correlations, either phenotypic or genetic, within male flowers of either species. However, there was a negative genetic correlation between ovary size and pistil number in females of T. dioicum and a negative phenotypic correlation between the number of pistils and the size of both stamens and pistils in females of T. pubescens. If negative correlations indicate the presence of trade-offs in allocation between traits, this agrees with Bateman s principle (1948), which asserts that female reproduction is more likely to be limited by resources, whereas male reproduction is more often limited by access to mates. There was also a negative phenotypic correlation between the number of pistils and the number of stamens within female flowers of T. pubescens. In other words, plants appear to be limited in the total number of reproductive organs that they can make within the flower: plants that make more pistils per flower also produce fewer stamens. This limitation in flower size may be due to either the amount of space or the number of primordial meristems within the flower and not just the amount of resources available. Pistils have three times the biomass as stamens (Davis SL, in preparation). However, despite their relatively small cost compared to pistils in terms of resources, producing more stamens reduces pistil production. 364 The Journal of Heredity 2001:92(4)

5 The genetic correlation between these traits mirrored the phenotypic correlation in sign, arguing that this may be a real trade-off, but was not statistically significant. The existence of negative genetic correlations may not be easily detected: for example, Mazer et al. (1999) found evidence of a negative genetic correlation between anther number and ovule number in Spergularia marina only after determining correlated responses to artificial selection. Correlations Between the Sexes Since the perianth in the two species studied here are relatively small compared to the reproductive structures within the flower, the size of the flowers in these species is most easily measured by the number of reproductive organs per flower. There was no genetic correlation between the number of pistils per flower in females and the number of stamens per flower in males in T. dioicum. Therefore the size of flowers in males and females should be able to respond independently to selection in this species. If the same genes control stamen production in males and females of T. pubescens, a positive genetic correlation should exist between these traits. Instead, the family-mean correlation between these traits was nonsignificant. On the other hand, the positive genetic correlation between stamen number in males and pistil number in females of T. pubescens still indicates that flower development in the two sexes is not independent, so selection acting differentially in each sex may not lead to separate evolutionary optima. Since pistils and stamens are not directly homologous, the correlation between these traits could be explained if it is flower size that is being genetically controlled, which would then determine the number of reproductive organs per flower that can be made. Furthermore, although the number of stamens in males is not directly correlated with the number of stamens in females, they are not independent. For example, families with large flowers would be comprised of males that have many stamens per flower and females that have many pistils per flower. The within-flower phenotypic correlations showed that females that produce many pistils per flower also produce fewer stamens. One possible explanation for the maintenance of the positive correlation between stamen number in males and pistil number in females would be if flower size was negatively correlated with flower number, so that families that have fewer flowers per plant also have bigger flowers in both males and females. Meagher (1994) found that artificial selection on flower size caused an indirect and inverse effect on flower number in Silene latifolia, suggesting a negative genetic correlation between these traits. A similar negative correlation between flower size and flower number may be maintaining the positive correlation between flower size in males and in females of T. pubescens. Since each pistil in females of T. pubescens is uniovulate, the number of pistils a female produces directly determines the number of seeds that female can potentially produce. If selection acts against stamen production in favor of more pistils, and hence more possible seeds, the negative correlation between stamen and pistil production within female flowers should be able to augment this selection and hasten a response toward further sexual dimorphism in T. pubescens. Two factors may help account for the maintenance of stamens. First, environmental variation may mask the trade-off between stamens and pistils in natural populations and reduce its effects ( Fry 1993; Houle 1991). Davis (in preparation) was able to detect a negative phenotypic correlation between these traits in only 1 of 4 years in the same natural population from which the seeds for this study were originally collected. This also happened to be a year when overall flower size was significantly smaller, suggesting reduced environmental variance in that year. So although there is a trade-off between stamen and pistil production, its effects on evolution may be limited. Second, conflicting selection pressures on flower size in males and flower size in females, along with the positive genetic correlation between these traits may prevent the loss of stamens in females. Males, whose reproduction is theoretically limited by access to mates, may be selected to have larger flowers with more stamens. Selection on increased flower size (more stamens) in males would result in a correlated response in females for more pistils. For females, whose reproduction may be limited by resources, producing more pistils leads to smaller pistils (as indicated by the negative phenotypic correlation), and hence possibly smaller, less viable seeds, so females may be under selection to produce flowers with fewer, larger pistils and more stamens. In this way, conflicting selection pressure in males and females, in conjunction with positive genetic correlations between males and females, may be maintaining genetic variation for stamen production in females. From the Department of Biology, Indiana University, Bloomington, Indiana. I wish to thank Lynda Delph, Edmund Brodie III, Harold Lindman, and Loren Rieseberg for guidance and comments on this work. I also wish to thank two anonymous reviewers for their comments. I especially want to thank D. J. McClellan, E. Levri, M. Levri, D. Dudle, and J. Walgenbach for help planting and transplanting Thalictrum. The greenhouse crew at Indiana University were important for their help in maintaining the plants. This work was supported through funds from the Indiana Academy of Sciences and the Department of Biology at Indiana University. Address correspondence to Sandra L. Davis, Department of Biology, University of Louisiana at Monroe, Monroe, LA 71203, or The American Genetic Association References Ashman TL, Quantitative genetics of floral traits in a gynodioecious wild strawberry, Fragaria virginiana; implications for the independent evolution of female and hermaphrodite floral phenotypes. Heredity 83: Bateman AJ, Intrasexual selection in Drosophila. Heredity 2: Bawa KS, Evolution of dioecy in flowering plants. Annu Rev Ecol Syst 11: Boivin B, American Thalictra and their Old World allies. Rhodora 46:377, , Campbell DR, Genetic correlation between biomass allocation to male and female functions in a natural population of Ipomopsis aggregata. Heredity 79: Charlesworth D and Charlesworth B, A model for the evolution of dioecy and gynodioecy. Am Nat 112: Charlesworth D and Morgan MT, Allocation of resources to sex functions in flowering plants. Philos Trans R Soc Lond 332: Charnov E, Maynard Smith J, and Bull J, Why be a hermaphrodite? Nature 263: Cheverud JM, Dow MM, and Leutenegger W, The quantitative assessment of phylogenetic constraints in comparative analysis: sexual dimorphism in body weight among primates. Evolution 39: Conner JK, Floral evolution in wild radish: the roles of pollinators, natural selection, and genetic correlations among traits. Int J Plant Sci 158:S108 S120. Conner JK and Via S, Patterns of phenotypic and genetic correlations among morphological and life-history traits in wild radish, Raphunus raphanistrum. Evolution 47: Cox PA, Niche partitioning between sexes of dioecious plants. Am Nat 117: Darwin C, On the different forms of flowers on plants of the same species. London: John Murray. Davis SL, Stamens are not maintained as attractants to pollinators in females of cryptically dioecious Thalictrum pubescens Pursch (Ranunculaceae). Sex Plant Reprod 10: Delph LF, Galloway LF, and Stanton ML, Sexual dimorphism in flower size. Am Nat 148: Elle E, Quantitative genetics of sex allocation in the andromonoecious perennial, Solanum carolinense (L.). Heredity 80: Fenster CB and Carr DE, Genetics of sex allocation in Mimulus (Scrophulariaceae). J Evol Biol 10: Downloaded from at Pennsylvania State University on March 6, 2014 Brief Communications 365

6 Fry J, The general vigor problem: can antagonistic pleiotropy be detected when genetic covariances are positive? Evolution 47: Geber MA, Fitness effects of sexual dimorphism in plants. Trends Ecol Evol 10: Houle D, Genetic covariance of fitness correlates: what genetic correlations are made of and why it matters. Evolution 45: Kaplan S and Mulcahy D, Mode of pollination and floral sexuality in Thalictrum. Evolution 25: Keener C, Studies in the Ranunculaceae of the southeastern United States. II. Thalictrum L. Rhodora 78: Lande R, Sexual dimorphism, sexual selection and adaptation in polygenic characters. Evolution 34: Lloyd DG and Webb CJ, Secondary sex characters in plants. Bot Rev 43: Mayer S and Charlesworth D, Cryptic dioecy in flowering plants. Trends Ecol Evol 6: Mazer SJ, Delesalle VA, and Neal PR, Responses of floral traits to selection on primary sexual investment in Spergularia marina: the battle between the sexes. Evolution 53: Mazer SJ and Hultgård UM, Variation and covariation among floral traits within and among four species of northern European Primula (Primulaceae). Am J Bot 80: Meagher TR, Sex determination in plants. In: Plant reproductive ecology: patterns and strategies (Lovett-Doust J and Lovett-Doust L, eds). New York: Oxford University Press; Meagher TR, The quantitative genetics of sexual dimorphism in Silene latifolia (Caryophyllaceae). I. Genetic variation. Evolution 46: Meagher TR, The quantitative genetics of sexual dimorphism in Silene latifolia (Caryophyllaceae). II. Response to sex-specific selection. Evolution 48: O Neil P and Schmitt P, Genetic constraints on the independent evolution of male and female reproductive characters in the tristylous plant Lythrum salicaria. Evolution 47: Rice WR, Analyzing tables of statistical tests. Evolution 43: Via S, The quantitative genetics of polyphagy in an insect herbivore. II. Genetic correlations in larval performance within and among host plants. Evolution 38: Willson MF, Sexual selection in plants. Am Nat 113: Received May 18, 2000 Accepted February 14, 2001 Corresponding Editor: William F. Tracy 366 The Journal of Heredity 2001:92(4)

Sexual reproduction in Angiosperms

Sexual reproduction in Angiosperms Sexual reproduction in Angiosperms Name: ANGIOSPERMS Angiosperms are plants that have their seeds enclosed in an ovule inside their flower. About 80% of the plants we see and know are angiosperms. The

More information

Dissect a Flower. Huntington Library, Art Collections, and Botanical Gardens

Dissect a Flower. Huntington Library, Art Collections, and Botanical Gardens Huntington Library, Art Collections, and Botanical Gardens Dissect a Flower Overview Students dissect an Alstroemeria or similar flower to familiarize themselves with the basic parts of a flower. They

More information

Biology 213 Angiosperms. Introduction

Biology 213 Angiosperms. Introduction Biology 213 Angiosperms Introduction The flowering plants, the angiosperms, are the most recent plants to evolve and quickly became the dominant plant life on this planet. They are also the most diverse

More information

not to be republished NCERT Heredity and Evolution CHAPTER 9 Multiple Choice Questions

not to be republished NCERT Heredity and Evolution CHAPTER 9 Multiple Choice Questions CHAPTER 9 Heredity and Evolution Multiple Choice Questions 1. Exchange of genetic material takes place in (a) vegetative reproduction (b) asexual reproduction (c) sexual reproduction (d) budding 2. Two

More information

Section 24 1 Reproduction With Cones and Flowers (pages 609 616)

Section 24 1 Reproduction With Cones and Flowers (pages 609 616) Chapter 24 Reproduction of Seed Plants Section 24 1 Reproduction With Cones and Flowers (pages 609 616) Key Concepts What are the reproductive structures of gymnosperms and angiosperms? How does pollination

More information

Pollination: Flower to Fruit

Pollination: Flower to Fruit Pollination: Flower to Fruit Answer Key Vocabulary: anther, cross pollination, filament, fruit, nectar, ovary, ovule, pedicel, petal, pistil, pollen, pollen tube, pollination, receptacle, self pollination,

More information

The Flower! What is the flower?

The Flower! What is the flower? The outstanding and most significant feature of the flowering plants (and that which sets them out from other vascular plants) is the flower. Understanding the flower structure and names of the parts is

More information

4th GRADE MINIMUM CONTENTS-NATURAL SCIENCE UNIT 11: PLANTS

4th GRADE MINIMUM CONTENTS-NATURAL SCIENCE UNIT 11: PLANTS PLANT BITS 4th GRADE MINIMUM CONTENTS-NATURAL SCIENCE UNIT 11: PLANTS There are four main parts to a plant. They are the root, stem, leaf and flower. Each part has an important task to do in the life of

More information

Dr. Sandra L. Davis December, 2013

Dr. Sandra L. Davis December, 2013 Department of Biology University of Indianapolis Indianapolis, IN 46227 Dr. Sandra L. Davis December, 2013 Education : Ph. D. in Biology, Indiana University, Bloomington, IN. : B.S. in Biology, Indiana

More information

Teacher packs in Experimental Science. Bio Pack 5. Examining flower structure

Teacher packs in Experimental Science. Bio Pack 5. Examining flower structure Teacher packs in Experimental Science Bio Pack 5 Examining flower structure Pack contents: A. Teachers Guide B. Students Guide C. Assessment Student s sheet D. Extensions to experiment E. Links to other

More information

Section 24 1 Reproduction With Cones and Flowers (pages 609 616)

Section 24 1 Reproduction With Cones and Flowers (pages 609 616) Chapter 24 Reproduction of Seed Plants Section 24 1 Reproduction With Cones and Flowers (pages 609 616) This section describes the reproductive structures of gymnosperms and angiosperms. It also explains

More information

Pollination Activity. Jennifer J. Weber and Laura B. Vary

Pollination Activity. Jennifer J. Weber and Laura B. Vary Pollination Activity Jennifer J. Weber and Laura B. Vary Instructions for Pollination Activity Supplies: To assemble plant envelopes 1.Make copies of slides #5-17, once for every 5 students 2.In the lower

More information

Expt. How do flowering plants do it without flagella? The journey to find an egg. What causes pollen grain germination and tube growth?

Expt. How do flowering plants do it without flagella? The journey to find an egg. What causes pollen grain germination and tube growth? 1 Expt. How do flowering plants do it without flagella? The journey to find an egg. What causes pollen grain germination and tube growth? File: F12-07_pollen Modified from E. Moctezuma & others for BSCI

More information

ISSECTING A LOWER Florida Sunshine State Standards Benchmark SC.F.1.3.1 Background Information:

ISSECTING A LOWER Florida Sunshine State Standards Benchmark SC.F.1.3.1 Background Information: ISSECTING A LOWER Florida Sunshine State Standards Benchmark SC.F.1.3.1 The student understands that living things are composed of major systems that function in reproduction, growth, maintenance, and

More information

Recommended Resources: The following resources may be useful in teaching this lesson:

Recommended Resources: The following resources may be useful in teaching this lesson: Unit A: Basic Principles of Plant Science with a focus on Field Crops Lesson 5: Understanding Flower Anatomy Student Learning Objectives: Instruction in this lesson should result in students achieving

More information

Plant Reproduction. 2. Evolutionarily, floral parts are modified A. stems B. leaves C. roots D. stolons E. suberins

Plant Reproduction. 2. Evolutionarily, floral parts are modified A. stems B. leaves C. roots D. stolons E. suberins Plant Reproduction 1. Angiosperms use temporary reproductive structures that are not present in any other group of plants. These structures are called A. cones B. carpels C. receptacles D. flowers E. seeds

More information

To meet the expectations of this unit, students should already know how the appearance of some organisms change over time.

To meet the expectations of this unit, students should already know how the appearance of some organisms change over time. GRADE 4: Life science 3 Life cycles of animals and plants UNIT 4L.3 12 hours About this unit This unit is the third of four units on life science for Grade 4. The unit is designed to guide your planning

More information

Chapter 2. Biology of Flowering Plants: Reproduction. Flowers and Pollination

Chapter 2. Biology of Flowering Plants: Reproduction. Flowers and Pollination BOT 3015L (Sherdan/Outlaw/Aghoram); Page 1 of 12 Chapter 2 Biology of Flowering Plants: Reproduction Flowers and Pollination Objectives Angiosperms. Understand the distinguishing characteristics of angiosperms.

More information

B.10B Interactions with Plants

B.10B Interactions with Plants B.10B Interactions with Plants Picture Vocabulary stimuli Anything that prompts a response or action response An action that is prompted by a stimulus xylem Plant tissue that transports water absorbed

More information

Parts of a Flower and Pollination

Parts of a Flower and Pollination Science Unit: Lesson 3: Soils, Plants, and First Nations Parts of a Flower and Pollination School year: 2007/2008 Developed for: Britannia Elementary School, Vancouver School District Developed by: Catriona

More information

The Evolution of Sex: Diversity

The Evolution of Sex: Diversity The Evolution of Sex: Diversity Lukas Schärer Evolutionary Biology Zoological Institute University of Basel 25.10.2011 Evolution of Sex and its Consequences HS 11 1 Two-fold cost of sex? 2 Summary sex

More information

Biology 172L General Biology Lab II Lab 03: Plant Life Cycles and Adaptations II: Gymnosperms and Angiosperms

Biology 172L General Biology Lab II Lab 03: Plant Life Cycles and Adaptations II: Gymnosperms and Angiosperms Biology 172L General Biology Lab II Lab 03: Plant Life Cycles and Adaptations II: Gymnosperms and Angiosperms Introduction Vascular seed-bearing plants, such as gymnosperms (cone-bearing plants) and angiosperms

More information

Chapter 38: Angiosperm Reproduction and Biotechnology

Chapter 38: Angiosperm Reproduction and Biotechnology Name Period Concept 38.1 Flowers, double fertilization, and fruits are unique features of the angiosperm life cycle This may be a good time for you to go back to Chapter 29 and review alternation of generation

More information

The Flower - what is it?! Floral structure will be examined in lab this Mon/Tues save space in your notes!

The Flower - what is it?! Floral structure will be examined in lab this Mon/Tues save space in your notes! The Flower - what is it?! Floral structure will be examined in lab this Mon/Tues save space in your notes! Magnoliophyta - Flowering Plants! Introduction to Angiosperms "angio-" = vessel; so "angiosperm"

More information

Exam 1. CSS/Hort 430. 2008 All questions worth 2 points

Exam 1. CSS/Hort 430. 2008 All questions worth 2 points Exam 1. CSS/Hort 430. 2008 All questions worth 2 points 1. A general definition of plants is they are eukaryotic, multi-cellular organisms and are usually photosynthetic. In this definition, eukaryotic

More information

What's in a Flower. Ages: 8 to 12. Contributor: Susan Jaquette, Cornell Plantations volunteer

What's in a Flower. Ages: 8 to 12. Contributor: Susan Jaquette, Cornell Plantations volunteer Ages: 8 to 12 What's in a Flower Contributor: Susan Jauette, Cornell Plantations volunteer Main idea: Flowers are composed of several distinct parts, each of which plays an important role in nature. Objective:

More information

DID YOU KNOW that the plants most important to

DID YOU KNOW that the plants most important to Flower Anatomy DID YOU KNOW that the plants most important to agriculture all produce flowers? Every major food crop is a flowering plant. We do not think about the flowers of wheat, rice, corn, and soybeans.

More information

Operation Flower Dissection

Operation Flower Dissection Operation Flower Dissection Classroom Activity: K-4 Time: One to two 50-minute class periods Overview: In this activity, students will observe the similarities and differences between flowers of different

More information

Introduction to genetics

Introduction to genetics Introduction to genetics Biology chapter 11 Mr. Hines 11.1 The work of Gregor Mendel What makes you unique? A. Nearly all living things are unique in some way. B. Humans for example all have different

More information

Microevolution: The mechanism of evolution

Microevolution: The mechanism of evolution Microevolution: The mechanism of evolution What is it that evolves? Not individual organisms Populations are the smallest units that evolve Population: members of a species (interbreeding individuals and

More information

STATION 1: Gymnosperm Survey

STATION 1: Gymnosperm Survey The Seed Plants: Laboratory Gymnosperms & Angiospserms 5 Introduction Gymnosperms and angiosperms are vascular, sporophyte-dominant plants that produce seeds. Although these heterosporous plants still

More information

CHAPTER 23 THE EVOLUTIONS OF POPULATIONS. Section B: Causes of Microevolution

CHAPTER 23 THE EVOLUTIONS OF POPULATIONS. Section B: Causes of Microevolution CHAPTER 23 THE EVOLUTIONS OF POPULATIONS Section B: Causes of Microevolution 1. Microevolution is generation-to-generation change in a population s allele frequencies 2. The two main causes of microevolution

More information

Dry Forest. Objectives Students will be able to draw and label a flower. Student will learn the importance of flowers and their anatomy.

Dry Forest. Objectives Students will be able to draw and label a flower. Student will learn the importance of flowers and their anatomy. Concepts Floral structure and importance of flowers HCPS III Benchmarks SC6.6.4 Duration 1 hour Dry Forest Source Material Digital Vocabulary pollinators whorls sepals petals stamen pollen anthers stigma

More information

Anthropology 160: The Human Life Course NM HED Area III: Laboratory Science Competencies UNM Core Area 3: Physical and Natural Sciences

Anthropology 160: The Human Life Course NM HED Area III: Laboratory Science Competencies UNM Core Area 3: Physical and Natural Sciences Anthropology 160: The Human Life Course NM HED Area III: Laboratory Science Competencies UNM Core Area 3: Physical and Natural Sciences Student Learning Objectives: 1. Define basic concepts in evolutionary

More information

Seed color gene with two alleles: R = purple (or red) allele (dominant allele) r = yellow (or white) allele (recessive allele)

Seed color gene with two alleles: R = purple (or red) allele (dominant allele) r = yellow (or white) allele (recessive allele) Patterns of Inheritance in Maize written by J. D. Hendrix Learning Objectives Upon completing the exercise, each student should be able to define the following terms gene, allele, genotype, phenotype,

More information

Exercise 7 Angiosperm Reproduction: Flowers and Fruits Biol 1012, S2008, Lee, Etterson, and Little

Exercise 7 Angiosperm Reproduction: Flowers and Fruits Biol 1012, S2008, Lee, Etterson, and Little Exercise 7 Angiosperm Reproduction: Flowers and Fruits Biol 1012, S2008, Lee, Etterson, and Little Goals Relate structures in a flower to the plant life cycle: alternation of generations. Identify floral,

More information

PLANT EVOLUTION DISPLAY Handout

PLANT EVOLUTION DISPLAY Handout PLANT EVOLUTION DISPLAY Handout Name: TA and Section time Welcome to UCSC Greenhouses. This sheet explains a few botanical facts about plant reproduction that will help you through the display and handout.

More information

Chapter 21 Active Reading Guide The Evolution of Populations

Chapter 21 Active Reading Guide The Evolution of Populations Name: Roksana Korbi AP Biology Chapter 21 Active Reading Guide The Evolution of Populations This chapter begins with the idea that we focused on as we closed Chapter 19: Individuals do not evolve! Populations

More information

THE FLOWER: PARTS OF THE FLOWER

THE FLOWER: PARTS OF THE FLOWER THE FLOWER: PARTS OF THE FLOWER Materials Large flower A flower for each child Presentation (This is usually a group presentation) 1. Say, I would like to talk about the parts of the flower. 2. Place a

More information

Name Period. 3. How many rounds of DNA replication and cell division occur during meiosis?

Name Period. 3. How many rounds of DNA replication and cell division occur during meiosis? Name Period GENERAL BIOLOGY Second Semester Study Guide Chapters 3, 4, 5, 6, 11, 14, 16, 17, 18 and 19. SEXUAL REPRODUCTION AND MEIOSIS 1. What is the purpose of meiosis? 2. Distinguish between diploid

More information

Name Date Hour Table #

Name Date Hour Table # Flower Structure Flowers vary in size, shape and color. The largest flower, with at diameter of one meter (three feet and three inches) and weighing about 9 kilograms (20 pounds), is born on a plant from

More information

INTRODUCTION TO GENETICS USING TOBACCO (Nicotiana tabacum) SEEDLINGS

INTRODUCTION TO GENETICS USING TOBACCO (Nicotiana tabacum) SEEDLINGS INTRODUCTION TO GENETICS USING TOBACCO (Nicotiana tabacum) SEEDLINGS By Dr. Susan Petro Based on a lab by Dr. Elaine Winshell Nicotiana tabacum Objectives To apply Mendel s Law of Segregation To use Punnett

More information

Non-Disjunction Review. tent/animations/content/mistakesmei osis/mistakesmeiosis.html

Non-Disjunction Review.  tent/animations/content/mistakesmei osis/mistakesmeiosis.html Non-Disjunction Review http://www.sumanasinc.com/webcon tent/animations/content/mistakesmei osis/mistakesmeiosis.html Lesson# 1.6- Genetic Diversity and Heredity Gregor Mendel (1822-1884) Pioneer of genetics

More information

Flower Model: Teacher Instructions Sepals Anther Stamens (male) Filament Stigma Pistil Style (female) Ovary Petals sepals petals stamens pistil

Flower Model: Teacher Instructions Sepals Anther Stamens (male) Filament Stigma Pistil Style (female) Ovary Petals sepals petals stamens pistil Flower Model: Teacher Instructions In order to better understand the reproductive cycle of a flower, take a look at some flowers and note the male and female parts. Most flowers are different; some have

More information

Effect of temperature on seed setting behaviour in rabi sorghum (Sorghum bicolor (L). Moench)

Effect of temperature on seed setting behaviour in rabi sorghum (Sorghum bicolor (L). Moench) Electronic Journal of Plant Breeding, 1(4): 776782 (July 2010) Research Article Effect of temperature on seed setting behaviour in rabi sorghum (Sorghum bicolor (L). Moench) anapati Mukri, B.D Biradar

More information

Diagram of a Typical Plant

Diagram of a Typical Plant Grade: 9 to 12 Length: variable Subjects: life science Topics: weed identification Objectives Exercises in this lesson help students achieve the following objectives: Understand the basic parts of a plant

More information

Petals Petals are designed to attract as many insects as possible to visit the flower. They have two main ways of doing this.

Petals Petals are designed to attract as many insects as possible to visit the flower. They have two main ways of doing this. Teacher s Fact File Learn: Flower There are many different types of flowers your class may find when walking through the park. Blues, yellows, purples and whites are usually the more common colours as

More information

Lec 2. Plant body: form & function

Lec 2. Plant body: form & function Lec 2. Plant body: form & function 1. Seed plants are the most successful land plants. Why? Main stages of a plant s life cycle. 2. Plants have developed appropriate structures to carry out the functions:

More information

Foundations of Genetics. Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display

Foundations of Genetics. Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display Foundations of Genetics 8.1 Mendel and the Garden Pea The tendency for traits to be passed from parent to offspring is called heredity Gregor Mendel (1822-1884) The first person to systematically study

More information

Meiosis and Life Cycles - 1

Meiosis and Life Cycles - 1 Meiosis and Life Cycles - 1 We have just finished looking at the process of mitosis, a process that produces cells genetically identical to the original cell. Mitosis ensures that each cell of an organism

More information

Achadiah Rachmawati Plant Callus

Achadiah Rachmawati Plant Callus Achadiah Rachmawati Plant Callus Callus, Cambium and Plant Reproduction Callus Definition Plant callus (plural calluses or calli) a mass of unorganized parenchyma cells derived from plant tissue (explants)

More information

Nature of Genetic Material. Nature of Genetic Material

Nature of Genetic Material. Nature of Genetic Material Core Category Nature of Genetic Material Nature of Genetic Material Core Concepts in Genetics (in bold)/example Learning Objectives How is DNA organized? Describe the types of DNA regions that do not encode

More information

The Toledo Zoo/ThinkingWorks. Teacher Overview for the Conservatory Lessons

The Toledo Zoo/ThinkingWorks. Teacher Overview for the Conservatory Lessons The Toledo Zoo/ThinkingWorks Teacher Overview for the Conservatory Lessons Teacher Overview: Conservatory Plants have many traits that are unique to this particular kingdom of living things. Below is a

More information

Curriculum Map: Life Sciences Grade 8. Month. Unit. Essential Questions (List 1-3 Essential Questions)

Curriculum Map: Life Sciences Grade 8. Month. Unit. Essential Questions (List 1-3 Essential Questions) Month Unit 2016-2017 Curriculum Map: Life Sciences Grade 8 September Scientific Inquiry October Origin of Life/Cellular Processes Essential Questions (List 1-3 Essential Questions) Learning Outcomes/Objectives

More information

Double Fertilization and Post - Fertilization Events: Measuring

Double Fertilization and Post - Fertilization Events: Measuring WFP062298 Double Fertilization and Post - Fertilization Events: Measuring Concepts In plants fertilization is the event in sexual reproduction which follows pollination. In higher plants, two sperm are

More information

Animal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri

Animal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri Animal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri Criteria for assessing the validity of animal models of human behavioral research Face validity:

More information

Mr. Storie 10F Science Reproduction Unit Review. Reproduction Review YOU ARE EXPECTED TO KNOW THE MEANING OF ALL THE FOLLOWING TERMS:

Mr. Storie 10F Science Reproduction Unit Review. Reproduction Review YOU ARE EXPECTED TO KNOW THE MEANING OF ALL THE FOLLOWING TERMS: Reproduction Review YOU ARE EXPECTED TO KNOW THE MEANING OF ALL THE FOLLOWING TERMS: CHROMOSOME GENE DNA TRAIT HEREDITY INTERPHASE MITOSIS CYTOKINESIS ASEXUAL BINARY FISSION CELL CYCLE GENETIC DIVERSITY

More information

Quiz #4 Ch. 4 Modern Evolutionary Theory

Quiz #4 Ch. 4 Modern Evolutionary Theory Physical Anthropology Summer 2014 Dr. Leanna Wolfe Quiz #4 Ch. 4 Modern Evolutionary Theory 1. T/F Evolution by natural selection works directly on individuals, transforming populations. 2. T/F A genotypic

More information

7 th Grade Science Genetics Review

7 th Grade Science Genetics Review 7 th Grade Science Genetics Review #1 The passing of traits from one generation to the next. A: Dominant traits B: Heredity C: Trait acquisition D: None of these B. Heredity #2 Which would result in the

More information

Plant Growth & Development. Growth Stages. Differences in the Developmental Mechanisms of Plants and Animals. Development

Plant Growth & Development. Growth Stages. Differences in the Developmental Mechanisms of Plants and Animals. Development Plant Growth & Development Plant body is unable to move. To survive and grow, plants must be able to alter its growth, development and physiology. Plants are able to produce complex, yet variable forms

More information

Modules 5: Behavior Genetics and Evolutionary Psychology

Modules 5: Behavior Genetics and Evolutionary Psychology Modules 5: Behavior Genetics and Evolutionary Psychology Source of similarities and differences Similarities with other people such as developing a languag, showing similar emotions, following similar

More information

Genetics: The Science of Heredity

Genetics: The Science of Heredity Chapter 3 Genetics: The Science of Heredity Objectives Describe the results of Mendel's Experiment. Identify the role of alleles in controlling the inheritance of traits. Page 70 This Baby Koala What is

More information

8. Study the cladogram underline the derived characteristics and circle the organisms that developed from them.

8. Study the cladogram underline the derived characteristics and circle the organisms that developed from them. Seed Plants: Gymnosperms and Angiosperms Answer the questions as you go through the power point, there are also paragraphs to read where you will need to hi-lite or underline as you read. 1. What are the

More information

AP BIOLOGY SUMMER WORK 2016

AP BIOLOGY SUMMER WORK 2016 AP BIOLOGY SUMMER WORK 2016 Welcome to AP Biology! AP Biology is a rigorous course designed to be equivalent to a first year biology course at a university. This summer work was designed to get you started

More information

A trait is a variation of a particular character (e.g. color, height). Traits are passed from parents to offspring through genes.

A trait is a variation of a particular character (e.g. color, height). Traits are passed from parents to offspring through genes. 1 Biology Chapter 10 Study Guide Trait A trait is a variation of a particular character (e.g. color, height). Traits are passed from parents to offspring through genes. Genes Genes are located on chromosomes

More information

Darwinian Natural Selection

Darwinian Natural Selection Darwinian Natural Selection Evidence of Evolution Direct observation: species change Fossils show intermediate forms Extant species show structural, developmental and genetic homology Vestigial traits

More information

Problem Set 4 BILD10 / Winter 2014

Problem Set 4 BILD10 / Winter 2014 1) The DNA in linear eukaryotic chromosomes is wrapped around proteins called, which keep the DNA from getting tangled and enable an orderly, tight, and efficient packing of the DNA inside the cell. A)

More information

Determining Acceptance of the 9:3:3:1 Ratio in Fruit Fly Crosses Using the Chi Squared Test

Determining Acceptance of the 9:3:3:1 Ratio in Fruit Fly Crosses Using the Chi Squared Test Determining Acceptance of the 9:3:3:1 Ratio in Fruit Fly Crosses Using the Chi Squared Test Abstract In this experiment we set out to determine whether or not two different fruit fly crosses fit the 9:3:3:1

More information

TEXAS A&M PLANT BREEDING BULLETIN

TEXAS A&M PLANT BREEDING BULLETIN TEXAS A&M PLANT BREEDING BULLETIN October 2015 Our Mission: Educate and develop Plant Breeders worldwide Our Vision: Alleviate hunger and poverty through genetic improvement of plants A group of 54 graduate

More information

NURSERY PROPAGULE COLLETION AND GROWING GUIDELINES TO AVOID GENETIC DEGRADATION ON RESTORATION SITES

NURSERY PROPAGULE COLLETION AND GROWING GUIDELINES TO AVOID GENETIC DEGRADATION ON RESTORATION SITES NURSERY PROPAGULE COLLETION AND GROWING GUIDELINES TO AVOID GENETIC DEGRADATION ON RESTORATION SITES GOLDEN GATE NATIONAL PARKS Betty L Young, Program Director of Nurseries In an ideal world, we would

More information

Programme Cycle Three

Programme Cycle Three Teachers Instructions Activity 1 Plants & Vegetation Plants can be either herbaceous or woody. Most Herbaceous Plants have stems that are soft, green, and contain little woody tissue. These plants are

More information

Chapter 9 Patterns of Inheritance

Chapter 9 Patterns of Inheritance Bio 100 Patterns of Inheritance 1 Chapter 9 Patterns of Inheritance Modern genetics began with Gregor Mendel s quantitative experiments with pea plants History of Heredity Blending theory of heredity -

More information

ALFALFA GENETIC MUTANTS. Edwin T. Bingham

ALFALFA GENETIC MUTANTS. Edwin T. Bingham ALFALFA GENETIC MUTANTS Edwin T. Bingham ebingham@wisc.edu Introduction Alfalfa mutants studied prior to 1967 are reported in USDA Bulletin No.1370 by D. K. Barnes. This report illustrates genetic mutants

More information

III.8. Evolutionary Limits and Constraints Ary Hoffmann. variation in characteristics inherited across generations

III.8. Evolutionary Limits and Constraints Ary Hoffmann. variation in characteristics inherited across generations III.8 Evolutionary Limits and Constraints Ary Hoffmann OUTLINE 1. Lack of genetic variation as a limit and constraint 2. Trade-offs 3. Multivariate selection 4. Gene flow in marginal populations limiting

More information

11.1 Traits. Studying traits CHAPTER 11. Breeds and traits. Genetics is the study of heredity

11.1 Traits. Studying traits CHAPTER 11. Breeds and traits. Genetics is the study of heredity 11.1 Traits Tyler has free earlobes like his father. His mother has attached earlobes. Why does Tyler have earlobes like his father? In this section you will learn about traits and how they are passed

More information

Genetic and Evolutionary Foundations of Behavior. Quick Question. Darwin s Theory 2/10/2012. Chapter 3

Genetic and Evolutionary Foundations of Behavior. Quick Question. Darwin s Theory 2/10/2012. Chapter 3 Genetic and Evolutionary Foundations of Behavior Chapter 3 Gray, Psychology, 6e Worth Publishers 2010 Quick Question What do you know about Darwin? Come up with as many things as possible. Darwin s Theory

More information

Photosynthesis happens inside the chloroplasts in a palisade cell like this one.

Photosynthesis happens inside the chloroplasts in a palisade cell like this one. 1.1 Plant organs Photosynthesis Photosynthesis is the way that plants make food. They use carbon dioxide and water to make glucose and oxygen. Photosynthesis is a chemical reaction. We can summarise it

More information

PLANT REPRODUCTION. Lesson Plans. Janet Huguet

PLANT REPRODUCTION. Lesson Plans. Janet Huguet Lesson Plans January March 2009 TOPIC: LIVING THINGS: HOW TO CLASSIFY THE TWO KINGDOMS LESSONS 1 & 2 TIMING 2 sessions LEVEL: KEY SKILLS: Pupils will be able To consider that scientific classification

More information

Angiosperms: Phylum Anthophyta, the flowering plants

Angiosperms: Phylum Anthophyta, the flowering plants Angiosperms: Phylum Anthophyta, the flowering plants 1. Overview of seed plant evolution Figure 29.7 Land plant evolution. 2. Traits of flowering plants a) Flowers b) Monocots vrs. Dicots 3. Pollination

More information

Meiosis and Sexual Life Cycles

Meiosis and Sexual Life Cycles Meiosis and Sexual Life Cycles Chapter 13 1 Ojectives Distinguish between the following terms: somatic cell and gamete; autosome and sex chromosomes; haploid and diploid. List the phases of meiosis I and

More information

2. For example, tall plant, round seed, violet flower, etc. are dominant characters in a pea plant.

2. For example, tall plant, round seed, violet flower, etc. are dominant characters in a pea plant. Principles of Inheritance and Variation Class 12 Chapter 5 Principles of Inheritance and Variation Exercise Solutions Exercise : Solutions of Questions on Page Number : 93 Q1 : Mention the advantages of

More information

Mendel suggested that flower colour was controlled by inherited factors. Draw a ring around the correct answer to complete the following sentences.

Mendel suggested that flower colour was controlled by inherited factors. Draw a ring around the correct answer to complete the following sentences. Q. The diagrams show one of Mendel s experiments. He bred pea plants. Mendel suggested that flower colour was controlled by inherited factors. Draw a ring around the correct answer to complete the following

More information

Quantitative genetics and reaction norms. What is quantitative genetics? How can a trait vary on a continuous scale?

Quantitative genetics and reaction norms. What is quantitative genetics? How can a trait vary on a continuous scale? Quantitative genetics and reaction norms Demography - analyses sources for variation in fitness Quantitative genetics - analyses the genetic reasons for variation in fitness How much genetic variation

More information

Sexual and Asexual Reproduction

Sexual and Asexual Reproduction Program Support Notes by: Janine Haeusler M. Sc (Ed), B. Ed Produced by: VEA Pty Ltd Commissioning Editor: Sandra Frerichs B.Ed, M.Ed. Executive Producers: Edwina Baden-Powell B.A, CVP. Sandra Frerichs

More information

Name: Verbal Reasoning. Science. Revision Guide. Sue Hunter AN HACHETTE UK COMPANY

Name: Verbal Reasoning. Science. Revision Guide. Sue Hunter AN HACHETTE UK COMPANY Name: Verbal Reasoning Science Revision Guide Sue Hunter AN HACHETTE UK COMPANY Contents and progress record Use this page to plot your revision. Colour in the boxes when you feel confident with the skill

More information

SEXUAL REPRODUCTION MRS. MAXEY

SEXUAL REPRODUCTION MRS. MAXEY SEXUAL REPRODUCTION MRS. MAXEY CHAPTER 1 SEXUAL REPRODUCTION As we discovered with asexual reproduction, not all living organisms reproduce asexually. So, now it is time to investigate the second type

More information

The Human Genome. Genetics and Personality. The Human Genome. The Human Genome 2/19/2009. Chapter 6. Controversy About Genes and Personality

The Human Genome. Genetics and Personality. The Human Genome. The Human Genome 2/19/2009. Chapter 6. Controversy About Genes and Personality The Human Genome Chapter 6 Genetics and Personality Genome refers to the complete set of genes that an organism possesses Human genome contains 30,000 80,000 genes on 23 pairs of chromosomes The Human

More information

JC BIOLOGY DEFINITIONS

JC BIOLOGY DEFINITIONS JC BIOLOGY DEFINITIONS 7 characteristics of living things - feeding, respiration, movement, sensitivity, growth, reproduction, excretion. Tissue similar cells doing the same job, muscle, blood, skin. Organ

More information

The Butterfly Project Pollinator Curriculum Guide

The Butterfly Project Pollinator Curriculum Guide The Butterfly Project Pollinator Curriculum Guide The Butterfly Project A Program of the Open Space Institute Chrissy Word, Project Coordinator Jill Weiss, Consultant The Butterfly Project Work Team the

More information

17. A testcross A.is used to determine if an organism that is displaying a recessive trait is heterozygous or homozygous for that trait. B.

17. A testcross A.is used to determine if an organism that is displaying a recessive trait is heterozygous or homozygous for that trait. B. ch04 Student: 1. Which of the following does not inactivate an X chromosome? A. Mammals B. Drosophila C. C. elegans D. Humans 2. Who originally identified a highly condensed structure in the interphase

More information

Evolution of Populations

Evolution of Populations Evolution of Populations Evolution Q: How can populations evolve to form new species? 17.1 How do genes make evolution possible? WHAT I KNOW SAMPLE ANSWER: There are different variations of the same gene.

More information

Practice Questions 1: Evolution

Practice Questions 1: Evolution Practice Questions 1: Evolution 1. Which concept is best illustrated in the flowchart below? A. natural selection B. genetic manipulation C. dynamic equilibrium D. material cycles 2. The diagram below

More information

CPO Science Life Science Grade Level 7

CPO Science Life Science Grade Level 7 CPO Science Life Science Grade Level 7 Louisiana Department of Education Science Correlation to Grade Level Expectations Grade 7 Correlation Document GRADE LEVEL EXPECTATIONS TO BE COMPLETED BY PUBLISHER

More information

1. chose the pea plant for 3 reasons: a. structure of the pea flower (more later) b. presence of distinctive traits c. rapid reproductive cycle 2.

1. chose the pea plant for 3 reasons: a. structure of the pea flower (more later) b. presence of distinctive traits c. rapid reproductive cycle 2. Genetics I. Genetics A. genetics: scientific study of heredity 1. we have known for centuries that traits are passed from parents to offspring 2. we didn t know how the traits were determined B. recall

More information

Genetic analysis on tiller number and plant height per plant in rice ( Or yza

Genetic analysis on tiller number and plant height per plant in rice ( Or yza ( ) 32 (5) : 527 534, 2006 Journal of Zhejiang University (Agric1 & Life Sci1) Article ID : 100829209 (2006) 0520527208 Genetic analysis on tiller number and plant height per plant in rice ( Or yza sativa

More information

Micropropagation of Date Palm (Phoenix Dactylifera)

Micropropagation of Date Palm (Phoenix Dactylifera) Micropropagation of Date Palm (Phoenix Dactylifera) What is plant propagation? The reproduction or increasing in number of plants. Can be done in one of two ways. Sexual. Asexual. Sexual Propagation Pollen

More information

Exam #2 BSC Fall. NAME Key answers in bold

Exam #2 BSC Fall. NAME Key answers in bold Exam #2 BSC 2011 2004 Fall NAME Key answers in bold _ FORM B Before you begin, please write your name and social security number on the computerized score sheet. Mark in the corresponding bubbles under

More information

COMPOST AND PLANT GROWTH EXPERIMENTS

COMPOST AND PLANT GROWTH EXPERIMENTS 6y COMPOST AND PLANT GROWTH EXPERIMENTS Up to this point, we have concentrated primarily on the processes involved in converting organic wastes to compost. But, in addition to being an environmentally

More information

TEST NAME: Genetics unit test TEST ID: GRADE:07 SUBJECT:Life and Physical Sciences TEST CATEGORY: School Assessment

TEST NAME: Genetics unit test TEST ID: GRADE:07 SUBJECT:Life and Physical Sciences TEST CATEGORY: School Assessment TEST NAME: Genetics unit test TEST ID: 437885 GRADE:07 SUBJECT:Life and Physical Sciences TEST CATEGORY: School Assessment Genetics unit test Page 1 of 12 Student: Class: Date: 1. There are four blood

More information

Assessment Schedule 2014 Biology: Demonstrate understanding of genetic variation and change (91157) Evidence Statement

Assessment Schedule 2014 Biology: Demonstrate understanding of genetic variation and change (91157) Evidence Statement NCEA Level 2 Biology (91157) 2014 page 1 of 5 Assessment Schedule 2014 Biology: Demonstrate understanding of genetic variation and change (91157) Evidence Statement NCEA Level 2 Biology (91157) 2014 page

More information