Phenology and reproductive effort of cultivated and wild forms of Pennisetum glaucum under experimental conditions in the Sahel : implications for the maintenance of polymorphism in the species
01 Jun 1996-Botany (NRC Research Press Ottawa, Canada)-Vol. 74, Iss: 6, pp 959-964
TL;DR: The wild and the cultivated forms of pearl millet, Pennisetum glaucum (L.) R.Br.
Abstract: In the Sahel region of Africa, the wild and the cultivated forms of pearl millet, Pennisetum glaucum (L.) R.Br., are sympatric and interfertile and yet have remained distinct for millenia. Reproductive barriers are not sufficient to explain this situation. To elucidate other possible mechanisms, the two forms were compared under experimental conditions in the Sahel for their phenology and reproductive effort. The length of the flowering period of each type was much longer than the average individual flowering period. When the last cultivated plants were finishing flowering, 65% of the wild plants were still flowering and 30% were just starting to flower. Thus, the last group was completely isolated from cultivated pearl millet gene flow (endogamic reproduction). The two forms of pearl millet also differed in the distribution of aboveground biomass among different plant parts, except for the number of seeds per plant. Both phenological behaviour and reproductive effort contribute to the maintenance of dist...
The seeds uscil liir tlic stiidy saiiiplcs wcrc collcctcd from hundreds of spikes.
The first D A E wiis the sanie for wild pearl niillet Lind cultiviited pearl millet, 3 days after sowing.
In riich pocket one randomly chosen plant wiis observed until the ciid of its cycle.
The llowcring phase for the cultivatctl li)riii hegins with tlic Il~i\vcriiig of the spike of the niain stcni (the lïrsi stciii ciiicrgcd) aiid crids with ;I spike of a priniary tiller.
I'llcnology or ~lowcrillg
HOWcvcr, the total Ilowcring tinic ol'thc siirill>lcs WU 45 d:iys l'or cultivated pc¿irl millet and iiiorc thcin 49 days for wild pcarl niillct, sincc the latter population had not finishcd Ilowcring at harvest (Fig. IO ).
The distribution over tirrie of thc pcrcentagc of' cultivatcd plants in thc flowering phasc followcd a noriilal curvc.
Flowcring bcgan at 42 DAE, whcri thc tillcring pliasc was finished.
Thc siinic typc of distribution was obscrvcd for the wild plunts.
Discussion
This study shows that undcr cxperinicnlal conditions, thc cultivatcd pcarl millct was always influenced by the wild pearl millet pollcnic cloud, whereas the wild pearl millct population escapcd from the cultivatcd pollenic cloud for a large part of its flowcring period.
The pared with those frolli dcviíint foriiis, Iikc sliibra.
TL;DR: A significantly lower number of alleles and lower gene diversity in cultivated pearl millet accessions than in wild accessions is shown, which contrasts with a previous study using iso-enzyme markers showing similar genetic diversity between cultivated and wild pearl Millet populations.
Abstract: Genetic diversity of crop species in sub-Sahelian Africa is still poorly documented. Among such crops, pearl millet is one of the most important staple species. In Niger, pearl millet covers more than 65% of the total cultivated area. Analyzing pearl millet genetic diversity, its origin and its dynamics is important for in situ and ex situ germplasm conservation and to increase knowledge useful for breeding programs. We developed new genetic markers and a high-throughput technique for the genetic analysis of pearl millet. Using 25 microsatellite markers, we analyzed genetic diversity in 46 wild and 421 cultivated accessions of pearl millet in Niger. We showed a significantly lower number of alleles and lower gene diversity in cultivated pearl millet accessions than in wild accessions. This result contrasts with a previous study using iso-enzyme markers showing similar genetic diversity between cultivated and wild pearl millet populations. We found a strong differentiation between the cultivated and wild groups in Niger. Analyses of introgressions between cultivated and wild accessions showed modest but statistically supported evidence of introgressions. Wild accessions in the central region of Niger showed introgressions of cultivated alleles. Accessions of cultivated pearl millet showed introgressions of wild alleles in the western, central, and eastern parts of Niger.
139 citations
Cites background from "Phenology and reproductive effort o..."
...Phenology (Renno and Winkel 1996), pollen competition (Sarr et al....
[...]
...Phenology (Renno and Winkel 1996), pollen competition (Sarr et al. 1988; Robert et al. 1991), and reproductive barriers (Amoukou and Marchais 1993) could explain this result....
TL;DR: It is found that a monophyletic origin of cultivated pearl millet in West Africa is the most likely scenario supported by the data set and the phylogenetic relationship among accessions not showing introgression is analyzed.
Abstract: During the last 12,000 years, different cultures around the world have domesticated cereal crops. Several studies investigated the evolutionary history and domestication of cereals such as wheat in the Middle East, rice in Asia or maize in America. The domestication process in Africa has led to the emergence of important cereal crops like pearl millet in Sahelian Africa. In this study, we used 27 microsatellite loci to analyze 84 wild accessions and 355 cultivated accessions originating from the whole pearl millet distribution area in Africa and Asia. We found significantly higher diversity in the wild pearl millet group. The cultivated pearl millet sample possessed 81% of the alleles and 83% of the genetic diversity of the wild pearl millet sample. Using Bayesian approaches, we identified intermediate genotypes between the cultivated and wild groups. We then analyzed the phylogenetic relationship among accessions not showing introgression and found that a monophyletic origin of cultivated pearl millet in West Africa is the most likely scenario supported by our data set.
TL;DR: Key examples of crop/wild sympatry and overlapping flowering phenology, pollen and seed dispersal, the barriers to hybridisation and introgression, the evolution and fate of interspecific hybrids, their fitness, and the potential cost of transgenes are reviewed.
Abstract: Crop-to-wild gene flow has received close attention over the past ten years in connection with the development and cultivation of transgenic crops. In this paper, we review key examples of crop/wild sympatry and overlapping flowering phenology, pollen and seed dispersal, the barriers to hybridisation and introgression, the evolution and fate of interspecific hybrids, their fitness, and the potential cost of transgenes. We pay particular attention to ways in which the evolution and divergence between crops and their wild relatives may interfere with these successive steps. Our review suggests that crop-to-weed gene flow is highly idiosyncratic and that crop gene dispersion will certainly be very difficult to preclude totally. Future directions for research should thus focus on the long-term establishment and effects of transgenes on natural communities.
105 citations
Cites background from "Phenology and reproductive effort o..."
...Conversely, overlaps can be limited, so that many wild plants will be isolated from the cultivated pollen source (Renno and Winkel, 1996)....
[...]
...overlaps can be limited, so that many wild plants will be isolated from the cultivated pollen source (Renno and Winkel, 1996)....
TL;DR: In this article, the authors assessed the impact of water deficits at three development stages: prior to flowering (S30), at the beginning of flowering and at the end of flowering in pearl millet grown in experimental conditions similar to Sahelian farming conditions.
Abstract: Several studies conducted under high input conditions have indicated little susceptibility of pearl millet to water deficit untill early grain filling, because the losses in main shoot production were fully compensated by increased tiller fertility. The present study assessed the impact of water deficits at three development stages: prior to flowering (S30), at the beginning of flowering (S45), and at the end of flowering (S60) in pearl millet grown in experimental conditions similar to Sahelian farming conditions. It included a control irrigation treatment simulating the natural distribution of rainfall throughout the cropping season. Both biomass production and grain yield were severely reduced by S30 and S45, while S60 had no effect. In S30 and S45, the flowering of tillers was delayed or totally inhibited. In both of these treatments, the low number of productive tillers did not compensate for damage to panicle initiation and flowering of the main shoot. All treatments maintained green leaves on the main shoot during the grain filling period, and in S30 leaf growth recovered from mid-season drought. These results illustrate how pearl millet mostly escapes drought by matching its phenology to the mean rainfall distribution in the Sahel. In the case of mid-season drought, some late productive tillers and the maintenance of green leaf biomass of the main shoots limited, but did not overcome, the yield losses. This study stresses the importance of agro-ecological conditions in control treatments, particularly the water regime and crop density, when assessing crop drought resistance.
TL;DR: In this article, the authors assessed the impact of water deficits at three develop- ment stages: prior to flowering, at the beginning of flowering, and at the end of flowering in pearl millet grown in experimental conditions similar to Sahelian farming conditions.
Abstract: l~lH~~~~il~~~~~l Abstract Several studies conducted under high input conditions have indicated little susceptibility of pearl millet to water deficit until1 early grain filling, because the Introduction losses in main shoot production were fully compens- ated by increased tiller fertility. The present study assessed the impact of water deficits at three develop- ment stages: prior to flowering (S30), at the beginning of flowering (545), and at the end of flowering (S60) in pearl millet grown in experimental conditions similar to Sahelian farming conditions. It included a control irrigation treatment simulating the natural distribution of rainfall throughout the cropping season. Both bio- mass production and grain yield were severely reduced by S30 and S45, while S60 had no effect. In S30 and S45, the flowering of tillers was delayed or totally inhibited. In both of these treatments, the low number of productive tillers did not compensate for damage The usual effects of drought on the development of a plant are a lowered production of biomass and/or a change in the distribution of this biomass among the different organs. The drought resistance of a cultivated plant reflects its capacity to limit the impact on the economic yield of these changes in biomass production and distribution. This capacity will depend on the devel- opment phase affected by water deficit, as well as on the intensity and the length of the drought. In Sahelian climates, where agrosystems are based essentially on pearl millet (Petznisetum gluucum (L.) R. Br.), the risks of drought can be classed into two types according to their impact on pearl millet cultivation. l i to panicle initiation and flowering of the main shoot. (1) Droughts at the beginning or end of the cropping All treatments maintained green leaves on the main season, frequent and often long, but to which peasant shoot during the grain filling period, and in S30 leaf farmers respond by successive re-sowing (in the case growth recovered from mid-season drought. These of early droughts) or by the choice of short-season results illustrate how pearl millet mostly escapes cultivars tolerant of post-floral water deficits (in the drought by matching its phenology to the mean rainfall case of late droughts); distribution in the Sahel. In the case of mid-season (2) Intermediate droughts, less frequent and often brief drought, some late productive tillers and the mainten- (about 10 d), but very unpredictable and without any ance of green leaf biomass of the main shoots limited, practical remedies. They affect development stages but did not overcome, the yield losses. This study such as initiation of panicles, earing, or flowering stresses the importance of agro-ecological conditions which, in cereals, are generally sensitive to water in control treatments, particularly the water regime deficits. The variability of these droughts in time and and crop density, when assessing crop drought space complicates the characterization of drought- __ .-- - resistant varieties. I l resistance.
"Phenology and reproductive effort o..." refers background in this paper
...The current wild form is distributed across the northern Sahel and is phylogenetically close to the ancestor of the cultivated pearl millet (Harlan 1975; Portbres 1976)....
TL;DR: An attempt is made to provide a framework in which both taxonomy and infraspecific classification can operate with a minimum of confusion.
Abstract: The methods of formal taxonomy have not been very satisfactory for the classification of cultivated plants. As a result, the people who deal with cultivated plants the most have developed their own informal and intuitive classifications based on experience as to what constitutes useful groupings. An attempt is made to provide a framework in which both systems can operate with a minimum of confusion. The structure of the total available gene pool is characterized by assigning taxa to primary, secondary and tertiary gene pools. At the infraspecific level, cultivars are grouped into races and subraces in an informal way without rigid rules for the use of terms.
TL;DR: Most modern interpreters of Cope's Rule have attributed its validity solely to certain fundamental advantages of size increase, at least one of which is alleged to have operated within most evolutionary lineages.
Abstract: Cope's Rule is the name customarily applied to the widespread tendency of animal groups to evolve toward larger physical size. According to Kurten (1953), this paleontologic rule of evolution "is second in repute only to 'Dollo's Law' of . . .irreversibility." Although the rule was never concisely formulated by Cope, it is generally implicit in his writings (Cope, 1887, 1896). Like many other evolutionary generalizations extracted from the fossil record, Cope's Rule was derived primarily from study of mammalian phylogeny. Writers like Deperet (1909), Newell (1949), and Rensch (1959) have added diverse examples of evolutionary size increase to the mammalian trends discussed by Cope. Because numerous exceptions are known, recognition of the concept as a law has been rejected by most workers. Still, it has been widely upheld as a valid empirical generalization, and of the definitions for "rule" listed by Webster, "a generally prevailing condition," describes it accurately. Most modern interpreters of Cope's Rule have attributed its validity solely to certain fundamental advantages of size increase, at least one of which is alleged to have operated within most evolutionary lineages (Newell, 1949, p. 122-123; Kurten, 1953, p. 105; Simpson, 1953, p. 151; Rensch, 1959, p. 210-211; Gould, 1966b, p. 1138). Most of the proposed advantages of evolutionary size increase have been reviewed by Newell (1949), Kurten (1953), Rensch (1959), and Gould (1966a). Among the more salient are: improved ability to capture prey or ward off predators, greater reproductive success, increased intelligence (with increased brain size), better stamina, expanded size range of acceptable food, decreased annual mortality, extended individual longevity, and increased heat retention per unit volume. Certainly when evolutionary size increase occurs, it is in response to selection pressure resulting from one or more advantages. As Bonner (1968) pointed out, however, a net trend toward increased size within a higher taxon usually produces an increase in mean and maximum animal size, but not necessarily in minimum animal size. Selection pressure favoring size decrease is not rare but only less common than pressure favoring size increase. Reliance solely on inherent advantages of larger size to explain the high incidence of size increase suffers from deficiencies resembling some of those attached to the discarded concept of orthogenesis. The "inherent advantage" idea attempts only to account for directional evolution, without reference to limits. Clearly when size increase occurs in a
TL;DR: The gene pool classification suggested by Harlan and de Wet (1971) is used in order to treat the several cereals on a uniform basis and is not intended to be a formal taxonomic system but rather a simple device to bring a genetic focus to bear on the taxonomies already available.
Abstract: First, we shall introduce the materials with which we are dealing, and this requires some understanding of formal names and botanical classification. The taxonomy of cultivated plants has long been in a state of confusion. The same array of variation is treated in radically different ways by different taxonomists (see Jirasek, 1966; Jeffrey, 1968). Classifications are cluttered with Latin names that have little or no biological meaning, and some individual taxa are given ranks ranging from variety to genus depending on who is doing the classifying. Inept classifications have probably caused more difficulty in understanding the origin and evolution of cultivated plants than any other factor. We shall use the gene pool classification suggested by Harlan and de Wet (1971) in order to treat the several cereals on a uniform basis. In this system the total array of variation within maximum genetic reach is partitioned into primary, secondary, and tertiary gene pools. The primary gene pool includes all those races that can be crossed with the crop, yielding reasonably fertile hybrids in which the chromosomes pair well and in whose offspring genetic segregation is reasonably normal. The primary gene pool corresponds to the widely accepted concept of the biological species. The secondary gene pool includes all those species that can be crossed with the crop but with restricted gene flow. Genes can be transferred from the secondary to the primary gene pool, but one must struggle with those barriers that separate biological species such as sterility, poor chromosome pairing, lethal or weak hybrids, or poorly adapted hybrid derivatives and so on. The tertiary gene pool includes all those species that can be crossed with the crop, but the hybrids lead essentially nowhere. The hybrids are lethal, completely sterile, or anomalous. If any gene transfer is possible at all, it must be through radical manipulation of some sort such as embryo culture, tissue culture, use of complex hybrid bridges and so on (see Harlan and de Wet, 1971). Polyploid series in cultivated plants pose some special problems. As a general rule, we have suggested (Harlan and de Wet, 1971) that each level be treated as a separate gene pool. The barriers between ploidy levels are not necessarily strong, however, and morphological differences are sometimes minimal and difficult to describe. Each series is different and appropriate treatments must be worked out crop by crop. Separate gene pools for ploidy levels in potato or sugarcane may not be appropriate at all. In the cereals considered here, the only problem of separation by ploidy level occurs in oats where A vena strigosa and A. barbata races are difficult to distinguish morphologically. Our gene pool classification is not intended to be a formal taxonomic system but rather a simple device to bring a genetic focus to bear on the taxonomies already available. The conventional epithets can be used without undue confusion pro-