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	<front>
		<journal-meta>
			<journal-id journal-id-type="publisher-id">pnut</journal-id>
			<journal-title-group>
				<journal-title>Peanut Science</journal-title>
			</journal-title-group>
			<issn pub-type="active">0095-3679</issn>
			<issn pub-type="ppub">0095-3679</issn>
			<publisher>
				<publisher-name>American Peanut Research and Education Society</publisher-name>
			</publisher>
		</journal-meta>
		<article-meta>
			<article-id pub-id-type="doi">10.3146/PS17-8.1</article-id>
			<article-id pub-id-type="sici">pnut-44-02-06</article-id>
			<article-id pub-id-type="publisher-id">PS17-8</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>ARTICLES</subject>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>Inheritance of Spear-Shaped Leaf in Peanut</article-title>
				<alt-title alt-title-type="runhead">Inheritance of Spear-Shaped Leaf</alt-title>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author">
					<string-name name-style="western">
						<given-names>William D.</given-names>
						<surname>Branch</surname></string-name>
					<xref rid="n101" ref-type="fn"><sup>1</sup></xref>
					<xref rid="cor1" ref-type="corresp"><sup>*</sup></xref>
				</contrib>
			</contrib-group>
			
			<pub-date pub-type="ppub">
				<month>7</month>
				<year>2017</year>
			</pub-date>
			<volume>44</volume>
			<issue>2</issue>
			<fpage>74</fpage>
			<lpage>76</lpage>
			<permissions><copyright-statement></copyright-statement>
				<copyright-year>2006</copyright-year>
			</permissions>
			<related-article related-article-type="pdf" xlink:href="PS17-8.1.pdf"></related-article>
			<abstract>
				<title>ABSTRACT</title>
				<p>Recently, a single Spear-shaped Leaf mutant plant was discovered in the 'Georgia-06G' peanut (<italic>Arachis hypogaea</italic> L. ssp. <italic>hypogaea</italic> var. <italic>hypogaea</italic>) cultivar. The mutant had narrow leaflets with each leaflet tapering to a point which gives the appearance of a spearhead shape. Three cross combinations were used to determine the inheritance of this new mutant. F<sub>1</sub>, F<sub>2</sub>, and F<sub>3</sub> segregation data strongly supported a single incompletely dominant gene, designated <italic>SpL</italic>, controlling the inheritance of the Spear-shaped Leaf trait. The F<sub>2:3</sub> homozygous spear-shaped individual plants had taller mainstem heights, narrower leaflet width, reduced pod weight, and lower SMK percentages compared to the F<sub>2:3</sub> homozygous normal leaf plants resulting from the same closely related cross combination (Georgia-06G x Spear-shaped Leaf mutant).</p>
			</abstract>
			<kwd-group>
				<title>Key Words</title>
				<kwd>groundnut</kwd><x xml:space="preserve">, </x>
				<kwd><italic>Arachis hypogaea</italic> L<x xml:space="preserve">.</x></kwd><x xml:space="preserve">, </x>
				<kwd>genetic ratios</kwd><x xml:space="preserve">, </x>
				<kwd>cross combinations</kwd>
			</kwd-group>
		</article-meta>
	</front>
	<body>
		<sec id="s1">
			<title>Introduction</title>
			<p>A small-leaf, spear-shaped leaflet peanut (<italic>Arachis hypogaea</italic> L.) mutant was first reported by <xref ref-type="bibr" rid="i0095-3679-44-2-74-Bhide1">Bhide and Desale (1970)</xref> and later by <xref ref-type="bibr" rid="i0095-3679-44-2-74-Desale1">Desale (1987)</xref>. This spear-shaped mutant was selected from a field increase of the semi-spreading cultivar 'Kopargaon No. 1' at Jalgaon, India during 1964. The mutant was subsequently crossed with Kopargaon No. 1, and the F<sub>1</sub> plants had normal leaves which suggest that this mutant was recessive. F<sub>2</sub> segregation fit a 3:1 ratio for normal and spear-shaped leaflet plants, respectively. The study by <xref ref-type="bibr" rid="i0095-3679-44-2-74-Bhide1">Bhide and Desale (1970)</xref>, showed that the small-leaf, spear-shaped mutant was controlled by recessive monogenic inheritance.</p>
			<p>During 2012, a new Spear-shaped Leaf mutant was isolated from the runner-type peanut cultivar 'Georgia-06G' (<xref ref-type="bibr" rid="i0095-3679-44-2-74-Branch2">Branch, 2007</xref>). This mutant likewise had small spear-shaped leaflets with each leaflet tapering to a point at the top and bottom and the widest section being in the middle which gives the appearance of a spearhead shape (<xref ref-type="fig" rid="i0095-3679-44-2-74-f01">Figure 1</xref>). The objective of this study was to determine the inheritance of the present Spear-shaped Leaf mutant.</p>
			<fig id="i0095-3679-44-2-74-f01" position="float" orientation="portrait">
				<label>Fig. 1.</label>
				<caption>
					<p>Shows a normal leaf shape (bottom) and the Spear-shape Leaf mutant (top) found in the runner-type peanut cultivar, Georgia-06G.</p>
				</caption>
				<graphic xlink:href="i0095-3679-44-2-74-f01.png" position="float" orientation="portrait"></graphic>
			</fig>
		</sec>
		<sec id="s2">
			<title>Materials and Methods</title>
			<p>Three cross combinations were made in the greenhouse between Georgia-06G x Spear-shaped Leaf, 'OLin' (<xref ref-type="bibr" rid="i0095-3679-44-2-74-Simpson1">Simpson <italic>et al.</italic>, 2003</xref>) x Spear-shaped Leaf, and 'Georgia-04S' (<xref ref-type="bibr" rid="i0095-3679-44-2-74-Branch1">Branch, 2005</xref>) x Spear-shaped Leaf during 2013. F<sub>1</sub>, F<sub>2</sub>, and F<sub>3</sub> populations were grown during 2014, 2015, and 2016, respectively.</p>
			<p>Each year, seed were space-planted 30.5 cm apart in two rows with variable length depending upon number of seed x 1.8 m wide beds on a Tifton loamy sand soil type (fine-loamy, siliceous, thermic, Plinthic Kandidult) at the agronomy research farm near the University of Georgia, Coastal Plain Experiment Station, Tifton Campus. Recommended cultural practices with irrigation were followed throughout the growing seasons.</p>
			<p>Individual plants were harvested near optimum maturity based upon number of days after planting and above-ground plant appearance. After harvest, peanut pods were dried with forced warm air to approximately 6% moisture content before weighing and shelling.</p>
			<p>Phenotypic classification was based on individual plants before digging and inverting. Segregation data was analyzed by chi-square analysis for goodness-of-fit (P≤0.05) to expected genetic ratios (<xref ref-type="bibr" rid="i0095-3679-44-2-74-Strickberger1">Strickberger, 1968</xref>). Least significant difference (LSD) t-test was used to compare the ten plant average between F<sub>2:3</sub> spear-shaped and normal leaf homozygous genotypes for mainstem height, leaflet length and width, pod weight, sound mature kernels (SMK), and 100 SMK weight.</p>
		</sec>
		<sec id="s3">
			<title>Results and Discussions</title>
			<p>Each F<sub>1</sub> plant from all three crosses had an intermediate Spear-shaped Leaf. This indicates that the Spear-shaped Leaf trait is incompletely dominant to normal leaves which is different from the first report by <xref ref-type="bibr" rid="i0095-3679-44-2-74-Bhide1">Bhide and Desale (1970)</xref>.</p>
			<p>The F<sub>2</sub> segregation from the two cross combinations (Georgia-06G x Spear-shaped Leaf and Georgia-04S x Spear-shaped Leaf) showed an acceptable fit for a 1 Spear-shape to 2 Intermediate to 1 Normal leaf expected ratio, respectively (<xref ref-type="table" rid="i0095-3679-44-2-74-t01">Table 1</xref>). However, the F<sub>2</sub> segregation from the OLin x Spear-shaped Leaf cross combination did not fit the 1:2:1 ratio. This deviation from the expected ratio could be due to several albino seedlings observed in this infraspecific cross between subspecies (ssp. <italic>fastigiata</italic> and ssp. <italic>hypogaea</italic>) which died shortly after emergence. Because of this infraspecific cross combination, the total chi-square value was also found to be significantly different at P≤0.05, however the summed and homogeneity chi-square values were each found acceptable for the 1:2:1 expected genetic ratio. These F<sub>2</sub> results also disagree with the earlier report for the small-leaf, spear-shaped leaflets (<xref ref-type="bibr" rid="i0095-3679-44-2-74-Bhide1">Bhide and Desale, 1970</xref>).</p>
			<table-wrap id="i0095-3679-44-2-74-t01" position="float" content-type="2col" orientation="portrait">
				<label>Table 1.</label>
				<caption>
					<p>F<sub>2</sub> plant segregation for number of spear, intermediate, and normal shaped leaf plants from three peanut cross combinations, 2015.</p>
				</caption>
				<graphic xlink:href="i0095-3679-44-2-74-t01.png" position="float" orientation="portrait"></graphic>
			</table-wrap>
			<p>F<sub>3</sub> results confirmed a single incomplete dominant inheritance model with a 1:2:1 expected ratio from F<sub>2:3</sub> heterozygous intermediate Spear-shaped Leaf plants (<sub>X</sub><sup>2</sup> = 2.400, P = 0.25 - 0.50). Likewise, the Spear-shaped Leaf plants and normal shaped leaf plants bred true-to-type in the F<sub>2:3</sub> population from Georgia-06G x Spear-shaped Leaf cross combination as was expected. These F<sub>2:3</sub> homozygous spear-shaped individual plants had taller mainstem, narrow leaflet width, reduced pod weight, and lower sound mature kernel (SMK) percentages compared to the F<sub>2:3</sub> homozygous normal leaf plants resulting from this same closely related cross combination at maturity (<xref ref-type="table" rid="i0095-3679-44-2-74-t02">Table 2</xref>). Early in the growing season, it also was observed that the spear-shaped F<sub>2:3</sub> plants appear to have more thrips damage than the normal plants.</p>
			<table-wrap id="i0095-3679-44-2-74-t02" position="float" content-type="2col" orientation="portrait">
				<label>Table 2.</label>
				<caption>
					<p>Ten plant average comparison between F<sub>2:3</sub> normal leaf vs spear-shaped peanut genotypes from the Georgia-06G x Spear-shaped Leaf cross combination, 2016.</p>
				</caption>
				<graphic xlink:href="i0095-3679-44-2-74-t02.png" position="float" orientation="portrait"></graphic>
			</table-wrap>
			<p>These findings strongly suggest that the inheritance of the present Spear-shaped Leaf mutant is controlled by a single incompletely dominant gene, designated (<italic>SpL</italic>). This data also disagrees with the past recessive monogenic inheritance for the spear-shaped leaflet trait reported by <xref ref-type="bibr" rid="i0095-3679-44-2-74-Bhide1">Bhide and Desale (1970)</xref> and <xref ref-type="bibr" rid="i0095-3679-44-2-74-Desale1">Desale (1987)</xref>. It would have been interesting to make allelism tests between these two spear-shaped mutants, unfortunately there were nearly 50 years between the occurrences of these two leaf mutants.</p>
		</sec>
	</body>
	<back>
		<ref-list>
			<title>Literature Cited</title>
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			<fn-group>
                <label><p><bold>Author Affiliations</bold></p></label>
				<fn id="n101" fn-type="current-aff">
					<label><sup>1</sup></label>
					<p>Professor, University of Georgia, Department of Crop and Soil Sciences, Coastal Plain Exp. Sta., 2360 Rainwater Road, Tifton, GA 31793-5766.</p>
				</fn>
				<corresp id="cor1">
					<label>*</label>Corresponding author email: <email>wdbranch@uga.edu</email>
				</corresp>
			</fn-group>
	</back>
</article>
