The domestication of Helianthus annuus L. (sunflower)

All modern domesticated sunflowers can be traced to a single center of domestication in the interior mid-latitudes of eastern North America. The sunflower achenes and kernels recovered from six eastern North American sites predating 3000 b.p. that document the early history of this important crop plant are reanalyzed, and two major difficulties in the interpretation of archaeological sunflower specimens are addressed. First, achenes and kernels obtained from a modern wild sunflower population included in a prior genetic study because of its minimal likelihood for crop-wild gene flow, and its close genetic relationship to domesticated sunflowers, provide a new and more tightly drawn basis of comparison for distinguishing between wild and domesticated achene and kernel specimens recovered from archaeological contexts. Second, achenes and kernels from this modern wild baseline population were carbonized, allowing a direct comparison between carbonized archaeological specimens and a carbonized modern wild reference class, thereby avoiding the need for the various problematic shrinkage correction conversion formulas that have been employed over the past half century. The need for further research on museum collections is underscored, and new research directions are identified.     

Morphometric Analysis of Sunflower (<Emphasis Type="Italic">Helianthus annuus</Emphasis> L.) Achenes from Mexico and Eastern North America<Superscript>1</Superscript>

Morphometric Analysis of Sunflower ( Helianthus annuus L.) Achenes from Mexico and Eastern North America. Sunflower (Helianthus annuus L.) has played a major role in the evolution of agricultural systems in the Americas. The discovery of ancient domesticated remains from archaeological deposits in pre-Columbian Mexico offers new dimensions to widely accepted viewpoints on the domestication pattern of H. annuus. Although American sunflower populations north of Mexico have been examined extensively, Mexican indigenous domesticated landraces have not been studied in any detail. In this study, we morphologically assessed wild and domesticated sunflower achenes from Mexico and compared them to similar datasets from eastern North America. Additionally, we evaluated the utility of four computer-assisted shape measurements in discriminating between wild and domesticated sunflower achenes (fruits) and compared variation in achene size among modern wild and cultivated populations from both Mexico and the U.S. We found that, of the shape parameters tested, none were informative in distinguishing wild achenes from domesticated varieties. Subsequent size analysis, using conventional parameters of length, width, and thickness, showed that modern wild populations from Mexico had smaller achenes compared to modern populations from eastern North America. Domesticated achenes unearthed from Mexican archaeological sites, however, were significantly larger than the early domesticated specimens recovered from eastern North America. Our methodological results indicate that variation in archaeological sunflower achenes is better described by conventional size parameters rather than computerized shape analysis.     

Morphological variability of achenes of some European populations of Lactuca serriola L.

This study focuses on the morphological variability of Lactuca serriola achenes in relation to eco-geographic features. Fifty L. serriola populations from four European countries, Czech Republic, Germany, the Netherlands and United Kingdom, were studied. Five morphological characters of the achenes - length and width of achene, length/width index, length of beak, and number of ribs - were evaluated. Significant differences exist in achene morphology of two leaf forms of L. serriola, forma serriola and forma integrifolia. Achenes of f. serriola are shorter, thinner, shorter beaked, lower length/width index, and higher number of ribs compared to f. integrifolia. There was significant variation in the measured characters. Statistical analysis indicated that achene length and width increased along an east-west transect from 2.95 to 3.35 mm and 0.93 to 1.00 mm, respectively. Mean beak length had a similar trend with the exception of German achenes. They had shorter beaks than achenes originating from the Czech Republic with 4.38 and Germany with 4.33 mm. The same trend was evident for L/W index from Czech with a ratio of 3.21 and Germany with 3.14. The number of ribs increased from east to the west in continental Europe, whereas the lowest number of ribs was recorded in achenes collected in Czech with 10.89 and the UK with 10.59. Achene morphology was significantly correlated with three eco-geographic features; longitude, latitude, soil texture of the habitats. The other eco-geographic factors, altitude and population size, did not significantly correlate with the studied characters of L. serriola achenes.     

Effects of nitrogen supply on growth, yield and yield components of safflower and sunflower

Safflower (Carthamus tinctorius L.) represents an important oil crop internationally and may have a certain production potential under low input conditions, particularly in organic farming systems, where the putatively low nutrient requirement would be an advantage. However, little is known about the nutrient requirements of safflower. This study was undertaken to determine the growth and yield response of safflower as compared to sunflower (Helianthus annuus L.) under different N supplies. Safflower and sunflower plants were grown in Mitscherlich pots containing equal volumes of sand, nutrient-poor limed soil, and perlite. Nitrogen supply was the same for both species (0.25, 0.5, 1.0, 1.5, and 2.0 g per pot) in the first year, but was raised for sunflower (0.5, 1.0, 2.0, 3.0, and 4.0 g per pot) in the second. Increased N supply enhanced plant growth and yield for both species. Growth and yield of safflower increased up to 1.0 g pot⁻¹, while the optimum for sunflower was 2.0 g pot⁻¹. Safflower out-yielded sunflower at low N supply, while at high N level the opposite occurred. Functional analysis according to Michaelis–Menten revealed that – in terms of yield formation – safflower is superior to sunflower under N-limited conditions. Safflower is generally more efficient than sunflower in concentrating N in their shoots. N concentration in the photosynthetically active youngest mature blade of both species exhibited the same functional relationship with the N supply, but safflower required a much lower leaf N concentration to produce optimal yield as compared to sunflower, indicating the higher efficiency of the former in terms of NUE. Yield components analysis revealed that in safflower, yield is tightly correlated with the number of capitula per plant and the mass per achene, both being strongly correlated, too. On the contrary, sunflower yield was merely determined by the number of achenes per capitulum, followed by the mass per achene. Path coefficient analysis showed that in safflower, the direct effects of the achene and leaf N content as well as the leaf dry matter on oil yield are small, and mediated principally via indirect effects on the number of achenes per capitulum, while for sunflower the number of achenes per capitulum exerts a strong direct effect. Dedicated to the late Prof. Dr. Burkhard Sattelmacher, who passed away on November 21, 2005.     

Upwind Movement of Achenes of Centaurea eriophora L. on the Ground

The lightly compressed achenes of Centaurea eriophora L. beara pappus composed of stiff bristles at their apex and have anelaiosome appendage at their base. The pappus is ineffectivein keeping the achene wind-borne but does serve to regulatethe movement of the achene on the ground in response to wind.In wind the achene swivels like a weather vane with the baseof the achene pointing into the wind. In weak wind the pappusbristles prevent the achene from blowing away. In stronger windthe bristles move due to their flattened, flexible, hinge-likebases and act like ratchets against the substratum, thus enablingthe achene to move upwind. In either case achenes remain ingroups. Ants are attracted to the elaiosome and disperse theachenes. Wind-induced movement was explored by testing acheneson various substrata in a wind tunnel at free-stream speedsbetween 2 and 7 m s^-1. Wind dispersal; Centaurea eriophora ; seeds; achenes     

Effects of pollen donors on seed formation inEspeletia schultzii (Compositae) populations at different altitudes

The influence of different pollen donors on seed formation was investigated in three populations ofEspeletia schultzii that differ in environmental conditions and life history characteristics. Self pollen and pollen from different donors (< 15m apart) within each population was used in a diallel design in order to test the genetic base of seed set variation. Three measures of seed formation were used: (1) achene number; (2) proportion of filled achenes (fruits) that distinguishes between achenes with seeds and empty achenes; (3) proportion of aborted seeds that distinguishes between viable and aborted seeds. Self-pollinations resulted in empty achenes. Achene number did not vary between the different pollen donors. A bimodal pattern of filled achenes was found in two populations in two consecutive years. On the other hand, a unimodal pattern was found in crosses between more distant donors (> 30m). These patterns seems to be the results of a sporophytic incompatibility system. Seed abortion was highest at the higher elevations and seems to be correlated with elevation rather than with any genetic effect.     

中国特有植物艾比湖沙拐枣(Calligonum ebi-nuricum)居群内果实性状的变异

以中国荒漠特有植物艾比湖沙拐枣(Calligonum ebi-nuricum)果实形态特征为研究对象,对分布于原产区的同一居群12个样株的果实进行形态特征比较研究。通过每株30粒果实的喙长、果实长度、果实宽度、果实形状、瘦果长度、瘦果宽度、瘦果形状、刺毛长度、头部刺毛长度、两肋间距、两刺间距11个性状指标的数值分析表明：在株内与株间均有变异,株间果实性状变异较株内果实性状变异大;代表果实和瘦果大小指标正相关性极显著（p＜0.01）;两肋间距与瘦果形状呈负相关（r=-0.297**,p＜0.01）。研究结果证明：其果实形状和瘦果形状（长度、宽度及长宽比）较为稳定,可作为艾比湖沙拐枣的分种依据,而其它果实性状可塑性较大,只能作为分类的参考。该研究结果可供沙拐枣属其它种的准确分类参考。     

The Relationship Between Achene Weight,Embryo Weight and Germination inTaraxacumApomicts

Apomicts provide the opportunity to investigate the variation in achene size and performance that is due to factors other than genetic differences between achenes. Achene characteristics of a single capitulum from each of 15 biotypes of apomicticTaraxacumwere investigated. Within a capitulum, achene fresh weight ranged from 4.45-fold to 13.74-fold. Overall, 59.8% of achenes germinated. The probability of any one achene germinating was influenced both by its source and by its weight and ranged from 0.049 to 0.902 between capitula and from 0.028 to 0.873 between weight classes. An investigation of the relationship between achene parts in three biotypes identified a linear trend for embryos to acquire a greater proportion of total achene weight as achene weight increases. Although the heaviest achenes showed the best germination, it is predicted that, for a given investment of maternal resources, seedling number would be maximized by producing achenes of a single intermediate weight.     

POPULATION DYNAMIC CONSEQUENCES AND EVOLUTION OF SEED TRAITS OF HETEROSPERMA PINNATUM (ASTERACEAE)

We summarize research on variation in achene morphology of Heterosperma pinnatum. Each flowering head has a broad range of achene morphologies. There is no between-year seed bank. There are no ecologically relevant differences in achene size, seedling size, or seedling growth rates of the different morphs. Achenes located centrally in the heads lose dormancy earlier during the period between autumn achene production and the onset (May-July) of the rainy season. This results in relatively more early emergence of central achenes. Awned central achenes also have greater adhesive dispersibility. Emergence, survival, and fecundity were measured for achenes sown into natural populations in central Mexico. When harsh conditions result in few seeds surviving to reproduce, the advantage tends to go to the more “conservative” peripheral achenes. Greenhouse experiments show a strong genetic component to the determination of achene proportions within and among populations and a variety of genetic correlations. Populations with a high proportion of awned achenes tend to be found in vegetation types where they are likely to be ephemeral. A high proportion of central achenes tend to be found in sites with little pre-rainy season precipitation. These patterns are interpreted in terms of selection for dispersal and the within-year timing of germination.     

Variability of the inflorescence ofMicroseris laciniata (Compositae: Cichorieae)

Quantitative characters of the flowering head of a garden population ofMicroseris laciniata were scored during the second, third, and fourth season of growth. Number of achenes per head, number of phyllaries per head and the average number of pappus parts per achene in single heads show significant plant to plant variation. Achenes per head and pappus parts per achene were scored in identical plants in two subsequent seasons. The number of pappus parts per achene varies freely between five and ten. This contrasts with annual species ofMicroseris in which either five or ten pappus parts are found, depending on the species. In spite of a clear plant-specific average of pappus parts, both high and low pappus part determination can be demonstrated in all specimens. The number of pappus parts depends on the position of an achene on the receptacle, marginal achenes usually having fewer pappus parts than central ones. This gradient is not closely correlated with the position of an achene on the genetic spiral.     

QTL analysis reveals different and independent modes of inheritance for diagnostic achene characters in Microseris (Asteraceae)

There are few reliable diagnostic morphological characters for species of the asteracean genus Microseris, and quantitative differences in the shapes of the achenes and the paleaceous pappus parts play a decisive role in species recognition. The genetic basis of species and strain differences in various characters has been studied previously, but little is known about quantitative characters of the achenes.

We performed a quantitative trait locus (QTL) analysis in the F2 of an interspecific cross between Microseris douglasii and M. bigelovii for achene length, achene diameter, achene shape, palea length, awn length, and achene pigmentation.

Independent inheritance of the main heritable achene characters – achene length, palea length and achene pigmentation – was revealed by the detection of distinct and specific QTLs for these characters. For palea length five QTLs with about equal phenotypic effects were mapped on four different linkage groups. Achene length and achene shape (achene length / achene diameter) were determined by two different genetic systems with one major gene and two modifiers. The detection of QTLs with a polarity of the effects opposite to that in the parental strains for achene length and achene shape reveals genetic variation for a potential increase in species differences. For the highly heritable trait, achene pigmentation, the bimodal F2 distribution suggested single-factor inheritance for absence versus presence of spots, with dominance for the spotted condition. However, only relatively weak QTL effects on that trait could be detected. Additional molecular markers (RAPDs, AFLPs) have to be tested for cosegregation with that major gene. The results are discussed in the context of different theories for the evolution of morphological characters.     

Phytochrome et germination des semences de Rumex

The Rumex alpinus L. achenes show a special type of positive photosensitivity: several red light irradiations are necessary to induce optimal germination. The achenes deprived of their pericarp lose their photosensitivity and germinate readily in the dark. By spectrophotometry in vivo, the presence of the Pr form of phytochrome is revealed in these seeds. Short periods at extreme temperatures (30°C or 5°C) induce a good proportion of achenes to germinate. The gibberellins are inefficient on achene germination contrary to benzyladenine which exhibits some activity. A slight improvement of GA₄ effects has been detected on scarified fruits.     

一年生短命植物疏齿千里光果实异形性的生态学意义

疏齿千里光(Senecio subdentatus)是分布在新疆北部古尔班通古特沙漠中的一种具异形瘦果的菊科一年生短命植物。将野外观测与室内实验相结合, 对该物种异形瘦果的形态、扩散和萌发特性, 以及异形瘦果产量与植株大小的关系进行了研究, 并对其生态学意义进行了探讨。结果表明: 疏齿千里光果序中的外围果和中央果均为柱形, 但前者为淡黄色, 后者为褐色, 且二者在大小、冠毛长度及果皮微形态等方面均存在明显差异。两种瘦果均以单个果实为扩散单元, 且在静止空气中的降落速度和在1 m·s^-1与2 m·s^-1风速下的扩散距离无显著差异, 说明虽然果实大小和冠毛长度对瘦果扩散具有不同的影响, 但对其整体扩散能力无明显影响。在各温变周期(5/2、15/2、20/10、25/15和30/15 ℃)处理中, 淡黄色外围果的萌发率均高于褐色中央果的, 且不同温度间两种瘦果的萌发率均存在显著差异, 但光照条件对其无显著影响。果序中的中央瘦果数明显多于外围果的, 且植株中外围果所占比例与植株大小间呈显著负相关关系, 而中央果所占比例与植株大小间呈显著正相关关系。这些特点说明, 该物种的小植株倾向于产生较多较易萌发的外围果, 大植株倾向于产生较多不易萌发的中央果。在古尔班通古特沙漠不可预测的极端环境中, 疏齿千里光可通过异形瘦果间的萌发差异及调节其不同大小植株中异形瘦果的比例, 来减少同胞后代之间的竞争, 增加其对不同微环境条件的生态适应性。     

Significance of achene characteristics and within-achene resource allocation in the germination strategy of tetraploid Aster pilosus var. pilosus (Asteraceae)

The interrelationships among achene weight, allocation to embryo and pericarp, and germination time were determined for 500 stratified achenes of tetraploid Aster pilosus Willd. var. pilosus. Only 52.6% of the achenes germinated. Germinated achenes were significantly heavier than ungerminated achenes. Germination time was independent of achene weight and embryo weight, but varied inversely with pericarp weight. Variable achene weight is evolutionarily advantageous. Heavy achenes are at an advantage in that their proportionately larger embryos and thinner pericarps facilitate germination, promoting competitive establishment of seedlings. Lighter achenes are also at an advantage through increased dispersibility, and their relatively thick pericarp provides a persistent seed bank. Evolutionary pressures presumably maintain the variability in achene weight of var. pilosus. These results are discussed in the context of the early midsuccessional ecology of var. pilosus.     

Effect of source–sink ratio on seed set and filling in sunflower (Helianthus annuus L.)

Poor seed development in sunflower may result from insufficient assimilate supply (source limitation). To test this hypothesis, the effects of changed source–sink ratio on seed set (measured as percentage of empty achenes) and seed filling (measured as dry mass per filled achene) in individual plants were investigated. Source–sink ratio, defined as leaf area per floret (LAF), was experimentally altered using invasive (floret removal, defoliation) and non‐invasive (pulse of chilling, short days or shading during leaf or floret initiation) treatments. Shading at floret initiation proved the most effective non‐invasive method. Generally, an increase, or decrease, in LAF improved, or impaired, both seed set and filling. Increasing LAF by 2.0 cm²[95% confidence interval (1.5, 2.5)] decreased the percentage of empty achenes by 36.9%‐points (?41.9, ?30.9) and increased dry mass per filled achene by 20.1 mg (13.6, 26.7) in the capitulum centre. The effect of source–sink ratio on seed set was always strongest in the centre, whereas peripheral whorls were not affected. Achene mass was affected in all parts of the capitulum. It is concluded that source limitation is a major cause for empty achenes in sunflower plants grown under non‐stress conditions.     

Temperature and oxygen regulation of oleate desaturation in developing sunflower (Helianthus annuus) seeds

The effect of low (10°C) and high (30°C) temperature on in vivo oleate desaturation has been studied in developing sunflower ( Helianthus annuus L.) seeds under conditions of different oxygen availability (capitulum, detached achenes or peeled seeds). In seeds remaining in the capitulum, only a part of the oleate newly synthesized at high temperature was desaturated to linoleate, whereas more oleate than that synthesized de novo was desaturated at low temperature. Achenes were only able to significantly desaturate oleate at low temperatures. In contrast, oleate desaturation was detected in peeled seeds incubated at low and high temperatures, showing the highest rate at 20°C. Hull removing dramatically increased the activity of the microsomal oleate desaturase (FAD2, EC 1.3.1.35) at all studied temperatures, although a long-term inactivation of the enzyme was observed at high temperatures. Low oxygen concentration (1–2%) obtained by respiration of peeled seeds incubated in sealed vials, brought about the inactivation of the enzyme. All these data suggest that temperature regulates oleate desaturation controlling the amount of oleate and the FAD2 activity. In addition, this enzyme seems to be also regulated by the availability of oxygen, which is affected inside the achene by its diffusion through the hull, and the competition with respiration, both factors being temperature-dependent.     

Genetic components of heterocarpy inMicroseris hybrid B87 (Asteraceae,Lactuceae)

Microseris B87 is derived from a single hybrid specimen betweenM. pygmaea with few, weakly hairy peripheral achenes and aM. bigelovii with many, strongly hairy peripheral achenes. Offspring through the F4 and F5 generations obtained by spontaneous selfing were analyzed for the segregation of quantitative and qualitative characters relating to achene dimorphism. The phenotypic effects of two previously identified unlinked genes determining the relative number of outer achenes are characterized in partially and completely homozygous sublines. We show that two morphological markers genetically linked to one of these genes are themselves regulated by the system inducing heterocarpy. Not more than two more unlinked genes are involved in the genetic basis of the heterocarpic response. The interaction of these genes in determining the heterocarpy phenotypes is discussed in the framework of a model postulating genes for a morphogen gradient across the capitulum and genes responding to this gradient.     

Inbreeding effects on fitness traits in the heterocarpic herb Leontodon autumnalis L. (Asteraceae)

Heterocarpic plants are characterized by the production of distinct types of fruits that usually differ in their ecological behavior. In the Asteraceae, differences are mainly found between peripheral non-dispersal and central dispersal achenes (single-seeded fruits). Inbreeding depression is considered as an evolutionary force as it may reduce several fitness traits, and in the case of heterocarpic plants, it could influence fitness traits (e.g., seed set, germination rate, growth rate) of each fruit morph, which may have important ecological and evolutionary consequences. In particular, differential effects on fitness traits and dispersal of selfed and outcrossed progeny can strongly determine the viability of extant populations and the potential to colonize new habitats. We conducted a hand-pollination experiment in greenhouse conditions to test whether inbreeding affects the fitness of achene morphs in the heterocarpic herb Leontodon autumnalis (Asteraceae). Results show that achene morphs significantly differ in their ecological behavior, peripheral achenes germinating more and faster than central achenes. The significant interaction between pollination treatment and achene morph for germination probability might indicate a link between dormancy and mating system in L. autumnalis: germination was higher for outcrossed achenes in central achenes whereas the opposite pattern was exhibited by peripheral achenes. Selfing dramatically reduced seed set, probably as a consequence of strong self-incompatibility mechanisms rather than inbreeding effects. Inbreeding depression significantly affected late life-cycle traits, such as growth rate and biomass at flowering. Overall, results suggest that inbreeding depression seems to be an important selective force maintaining outcrossing in L. autumnalis.     

DIFFERENTIAL GERMINATION OF RAY AND DISC ACHENES IN HEMIZONIA INCRESCENS (ASTERACEAE)

The differential germination responses of ray and disc achenes of Hemizonia increscens (Asteraceae) were compared in field and laboratory investigations in order to gain insight into the ecological and evolutionary significance of heterocarpy. In the field, 200 ray and 200 disc achenes were placed in native, sterilized soil in a series of cleared, randomized, replicated plots. In a nearby plot a similar number of achenes were placed in plastic petri dishes in which high moisture conditions were maintained. Disc achene germination occurred under relatively minimal moisture conditions (<1 cm rainfall for 19 days) and relatively mild temperature regimes (21–7 C). Disc achene germination began three days after planting in the field plot and four days after they were put into the field petri dishes. In contrast, the onset of ray achene germination occurred 21 days after planting in the field plot and 19 days after planting in the field petri dishes. Averages of 2.05 and 2.71 disc achenes/day germinated in the field plot and field petri dishes, respectively. These contrasted with averages of 0.57 and 0.50 ray achenes/day germinated in the field plot and in the field petri dishes, respectively. A total of 67.5% and 69% disc achenes germinated in the field plot and the field petri dishes, but only 18% and 16.5% ray achenes germinated in the field plot and field petri dishes, respectively. Three separate treatments, using 100 ray and 100 disc achenes in each, were performed in laboratory growth chambers: 1) nicking the fruit coat, 2) excising the embryo, and 3) leaving the fruit coat intact. Onset of germination for all disc achene treatments occurred after three days. No significant differences were found among the three disc achene treatments in timing, rate, or germination percentage. All three disc treatments in the laboratory closely paralleled those for disc achenes in the field plots in time and germination percentage, but rates of germination were not as high. Germination of the nicked and excised ray achenes treatments began after four days, while germination of the untreated ray achenes began after 27 days. Untreated ray achenes in the laboratory paralleled the ray achenes in both field experiments in rate and germination percentage, but were delayed in time of germination. The nicked and excised ray achene treatments, however, were similar to the disc achene treatments in time of germination, and were not significantly different from disc achenes in rate of germination. These data suggest 1) that ray and disc achenes are markedly different in germination under identical conditions in field and laboratory experiments, and 2) this difference in germination response may be due to the thicker pericarp of the ray achenes.