--- title: "Heterosis, maternal effects and Gardner and Eberhart" author: "Muhammad Yaseen" date: last-modified format: html: self-contained: true number-sections: true toc: true toc-depth: 3 highlight-style: pygments code-tools: true html-math-method: mathjax vignette: > %\VignetteIndexEntry{Heterosis, maternal effects and Gardner and Eberhart} %\VignetteEngine{quarto::html} %\VignetteEncoding{UTF-8} execute: echo: true warning: false message: false --- {{< include _DiallelAnalysisR-macros.txt >}} ```{r setup, include = FALSE} knitr::opts_chunk$set(collapse = FALSE, comment = "", fig.width = 7, fig.height = 4.2, dev = if (knitr::is_latex_output()) "cairo_pdf" else if (requireNamespace("svglite", quietly = TRUE)) "svglite" else "png") if (requireNamespace("DiallelAnalysisR", quietly = TRUE)) library(DiallelAnalysisR) else pkgload::load_all(export_all = FALSE, helpers = FALSE, quiet = TRUE) ``` # Abstract When the parents are grown beside their crosses, the crosses can be compared with the parents: the heterosis of each cross, and Gardner and Eberhart's split of the crosses into variety effects and heterosis. In a diallel with reciprocals, the reciprocal effects split into maternal and non-maternal parts. This vignette takes these three in turn. # Heterosis With the parents grown beside the crosses, `Heterosis()` gives for each cross the heterosis over the mid-parent $MP = (P_i + P_j)/2$, the better parent and, if a check variety is named, the standard variety, in percent, each with a $t$ test. With $B$ replicates the standard errors of the differences are $\sqrt{3MS_E/(2B)}$ for $F_{ij} - MP$ and $\sqrt{2MS_E/B}$ for the other two. ```{r het} het <- Heterosis( y = Yield , Rep = Rep , Cross1 = Cross1 , Cross2 = Cross2 , data = GriffingData2 , Check = 3 , Better = c("higher", "lower")[1] ) het$StdErr head(x = het$Heterosis[order(-het$Heterosis$MP)], n = 6) ``` ```{r het-plot, fig.height = 6} plot(x = het, which = c("MP", "BP", "SH", "residuals", "qq")[1]) ``` For traits where a lower value is better (days to flowering, disease score), `Better = "lower"` takes the lower parent as the better one. # Maternal and non-maternal reciprocal effects Cockerham and Weir (1977) write the reciprocal effect of Methods 1 and 3 as $r_{ij} = m_i - m_j + n_{ij}$: a maternal effect of each parent and a non-maternal remainder. `Griffing(..., Reciprocal = "split")` splits the reciprocal line of the analysis of variance accordingly, on $P - 1$ and $(P - 1)(P - 2)/2$ degrees of freedom: ```{r recip} rs <- Griffing( y = Yield , Rep = Rep , Cross1 = Cross1 , Cross2 = Cross2 , data = GriffingData1 , Method = 1 , Model = 1 , Reciprocal = c("combined", "split")[2] ) rs$ANOVA round(x = rs$Maternal, digits = 3) round(x = rs$StdErr[c("mi", "mi_mj", "nij")], digits = 3) ``` Here neither part is significant: the reciprocal differences of this diallel are small and show no maternal pattern. # Gardner and Eberhart's Analyses II and III For parents and one set of crosses, Gardner and Eberhart (1966) model the parents by variety effects $v_j$ and the crosses by the mean of their parents plus heterosis. Analysis II splits the heterosis into average heterosis $\bar{h}$, variety heterosis $h_j$ and specific heterosis $s_{jj'}$: ```{r ge2} ge2 <- GardnerEberhart( y = Yield , Rep = Rep , Cross1 = Cross1 , Cross2 = Cross2 , data = GriffingData2 , Analysis = 2 , Model = 1 ) ge2$ANOVA ge2$Average ``` ```{r ge2-plot} plot(x = ge2, which = c("effects", "specific", "residuals", "qq")[1]) ``` Analysis III keeps the variety effects for the parents and analyses the crosses by general and specific combining ability, with average heterosis between the two; its gca and sca lines are Griffing's Method 4: ```{r ge3} ge3 <- GardnerEberhart( y = Yield , Rep = Rep , Cross1 = Cross1 , Cross2 = Cross2 , data = GriffingData2 , Analysis = 3 , Model = 1 ) ge3$ANOVA ``` # Summary `Heterosis()` compares each cross with its parents and a check; `Griffing(..., Reciprocal = "split")` asks whether the reciprocal differences come from the mother; `GardnerEberhart()` describes parents and crosses with one model of variety effects and heterosis. # References Cockerham, C. C. and Weir, B. S. (1977). Quadratic analyses of reciprocal crosses. *Biometrics*, 33(1), 187--203. Gardner, C. O. and Eberhart, S. A. (1966). Analysis and interpretation of the variety cross diallel and related populations. *Biometrics*, 22(3), 439--452. Murray, L. W., Ray, I. M., Dong, H. and Segovia-Lerma, A. (2003). Clarification and reevaluation of population-based diallel analyses: Gardner and Eberhart Analyses II and III revisited. *Crop Science*, 43(6), 1930--1937. Singh, R. K. and Chaudhary, B. D. (2004). *Biometrical Methods in Quantitative Genetic Analysis*. Kalyani Publishers, New Delhi.