Mathematics inspired by Darwin. Adaptive dynamics of dispersal and cooperation

In 1859, Charles Darwin published his theory of evolution by natural selection, the process occurring based on fitness benefits and fitness costs at the individual level. Traditionally, evolution has been investigated by biologists, but it has induced mathematical approaches, too. For example, adaptive dynamics has proven to be a very applicable framework to the purpose. Its core concept is the invasion fitness, the sign of which tells whether a mutant phenotype can invade the prevalent phenotype. In this thesis, four real-world applications on evolutionary questions are provided. Inspiration for the first two studies arose from a cold-adapted species, American pika. First, it is studied how the global climate change may affect the evolution of dispersal and viability of pika metapopulations. Based on the results gained here, it is shown that the evolution of dispersal can result in extinction and indeed, evolution of dispersalshould be incorporated into the viability analysis of species living in fragmented habitats. The second study is focused on the evolution of densitydependent dispersal in metapopulations with small habitat patches. It resulted a very surprising unintuitive evolutionary phenomenon, how a non-monotone density-dependent dispersal may evolve. Cooperation is surprisingly common in many levels of life, despite of its obvious vulnerability to selfish cheating. This motivated two applications. First, it is shown that density-dependent cooperative investment can evolve to have a qualitatively different, monotone or non-monotone, form depending on modelling details. The last study investigates the evolution of investing into two public-goods resources. The results suggest one general path by which labour division can arise via evolutionary branching. In addition to applications, two novel methodological derivations of fitness measures in structured metapopulations are given.

Response of weed populations to tillage, reduced herbicide and fertilizer rates in wheat (Triticum aestivum) production

Field experiments were carried out in 1999 and 2000 to investigate the effects of conventional (CT) and no-tillage (NT) systems, interacting with three herbicide dose levels and three nitrogen (N) levels on weed growth and wheat production of two varieties. There was a higher grain yield for NT system compared with CT in one year. CT weed biomass was lower than from NT weed biomass, in both varieties. No differences on wheat biomass and grain yield were observed between full and reduced herbicide rates. N fertilizer increased wheat biomass and grain yield significantly. Only N medium level had an effect upon weed biomass with respect to non-fertilized plots, while the highest fertilization rate lowered weed biomass. Conventional tillage, reduced herbicide rates and nitrogen fertilization were effective ways of limiting weed production in wheat.

Weed control and green ear yield in maize

The objective of this work was to evaluate the influence of weeding frequency on cultivar Centralmex green corn yield. Two experiments were conducted in Mossoró-RN (Brazil), with the use of sprinkler irrigation. A random block design with four replicates was used. It was observed that the total number and weight (TW) of unhusked green ears, the number and weight of marketable unhusked ears and the number and weight of marketable husked ears were reduced under no weeding treatment. The number timing of weedings did not influence green corn yield, except for one weeding at 60 DAP, which was equivalent to the "no weeding" treatment, for TW. When maize is marketed considering the total number of green ears, higher net income is obtained when one weeding is carried out 45 days after planting.

Weed control in young coffee plantations through post emergence herbicide application onto total area

This study was carried out to investigate the efficiency of several herbicides under field conditions, by post-emergence application onto the entire area, their effect on the control of weeds in young coffee plantations and commercial coffee and bean intercropping system, as well as on both crops. Seedlings of Coffea arabica cv. Red Catuaí with four to six leaf pairs were transplanted to the field and treated according to conventional agronomic practices. A bean and coffee intercropping system was established by sowing three lines of beans in the coffee inter-rows. At the time the herbicides were sprayed, the coffee plants had six to ten leaf pairs; the bean plants, three leaflets; and the weeds were at an early development stage. Fluazifop-p-butyl and clethodim were selective for coffee plants and controlled only Brachiaria plantaginea and Digitaria horizontalis efficiently. Broad-leaved weeds (Amaranthus retroflexus, Bidens pilosa, Coronopus didymus, Emilia sonchifolia, Galinsoga parviflora, Ipomoea grandifolia, Lepidium virginicum, and Raphanus raphanistrum) were controlled with high efficiency by sole applications of fomesafen, flazasulfuron, and oxyfluorfen, except B. pilosa, C. didymus, and R. raphanistrum for oxyfluorfen. Sequential applications in seven-day intervals of fomesafen + fluazifop-p-butyl, or clethodim, and two commercial mixtures of fomesafen + fluazifop-p-butyl simultaneously controlled both types of weed. Cyperus rotundus was only controlled by flazasulfuron. Except for fluazifop-p-butyl and clethodim, all herbicide treatments caused only slight injuries on younger coffee leaves. However, further plant growth was not impaired and coffee plant height and stem diameter were therefore similar in the treatments, as evaluated four months later. Fomesafen, fluazifop-p-butyl, and clethodim, at sole or sequential application, and the commercial mixtures of fomesafen + fluazifop-p-butyl were also highly selective for bean crop; thus at doses recommended for bean crop, these herbicides may be applied to control weeds in coffee and bean intercropping systems by spraying the entire area.

Effect of plant row spacing and herbicide use on weed aboveground biomass and corn grain yield

The use of narrow plant spacing in corn (Zea mays) has been suggested as a technological alternative to obtain grain yield increases, due to a better use of resources. The regular pattern could diminish intraspecific competition while favoring interspecific competition with weeds. The objective of this study was to analyze the effect of corn row spacing on weed aboveground biomass and corn grain yield. Field experiments were conducted during 2002/2003 and 2003/2004 growing seasons. Three corn hybrids with two-row width (0.70 and 0.35 m) were tested. A greater photosynthetic photon flux density (PPFD) interception with a lower weed aboveground dry matter in narrow row arrangement was obtained. Corn grain yield was greater in the narrow row arrangement than in the wide row spacing. This increase in grain yield was related to a better resource use that allows for a reduced interspecific competition. The use of reduced spatial arrangement appeared to be an interesting alternative to increase both the grain yield potential and corn suppressive ability against weeds in corn dryland production systems.

Effects of weed species competition on the growth of young coffee plants

The effects of competition of seven weed species on the growth of coffee plants were evaluated under greenhouse conditions. Thirty days after coffee seedling transplantation into 12 L pots with soil level area of 6.5 dm², weeds were transplanted into or sown in those pots, at densities of 0, 1, 2, 3, 4 and 5 plants per pot. Competition or weedy periods from weed transplantation or emergence to plant harvesting, at weed pre-flowering stage, were: 77 days - Bidens pilosa, 98 days - Brachiaria decumbens, 180 days - Commelina diffusa, 82 days - Leonurus sibiricus, 68 days - Nicandra physaloides, 148 days - Richardia brasiliensis and 133 days - Sida rhombifolia. Coffee plant height, stem diameter, leaf number and shoot dry matter were determined. Effects of competition by N. physaloides and S. rhombifolia against coffee plants were among the lowest, since only a slight decrease in all the characteristics evaluated in coffee plants was observed. The other weed species caused severe decrease in growth, mainly with increasing weed plant densities. Competition degree was found to depend on weed species and density.

Effects of weed control through cowpea intercropping on mayze morphology and yield

Intercropping combined with competitive maize cultivars can reduce the use of herbicides to control weeds. The objective of this work was to evaluate the effects of intercropping cowpea and maize, as well as hand-weeding on maize morphology and yield. The experimental design was in randomized complete blocks, with treatments arranged in split-plots and five replications. The plots consisted of four maize cultivars (BA 8512, BA 9012, EX 4001, EX 6004) and the split-plots consisted of the following treatments: no-weeding; twice hand-weeding (20 and 40 days after sowing); and intercropping with cowpea ('Sempre Verde' cultivar), both maize and cowpea sown at the same time. The variables evaluated were: maize fresh green ears and grain yield; characteristics of internodes, leaves, tassels, ears, grains; plant height and ear insertion height; number of weed plants and species; fresh and dry biomass of weed species and cowpea. Ten weed species were outstanding during the experiment, many of them from the Poaceae family. No interactions were found between weed control method and maize cultivars for most variables evaluated; and plants from hand-weeded split-plots showed superior mean values compared to plants from non-weeded and intercropped split-plots, both not differing from each other. The cowpea was inefficient in controlling weed, reducing the maize yields and not producing any grain. The maize cultivars 'BA 8512' and 'BA 9012 showed the highest mean green ear yield, and the highest grain yield in hand-weeded, no-weeded and intercropped split-plots. On the other hand, the maize cultivar 'EX 6004' showed such high means only in no-weeded and intercropped split-plots. 'EX 4001 presented the worst means in these variables for hand-weeded, no-weeded ant intercropped split-plots.

Growth and nutrient concentration in coffee root system under weed species competition

The effects of competition of six weed species on growth, nutrient concentration and nutrient content of coffee plant root system under greenhouse conditions were evaluated. Thirty days after coffee seedling transplantation into 12 L pots with soil level area of 6.5 dm². Weeds were transplanted or sowed in these pots, at densities of 0, 1, 2, 3, 4 and 5 plants per pot. The duration of competition (or weedy periods) from weed transplantation or emergence until plant harvesting, at the weed preflowering stage, were (in days): 77 (Bidens pilosa), 180 (Commelina diffusa), 82 (Leonurus sibiricus), 68 (Nicandra physaloides), 148 (Richardia brasiliensis) and 133 (Sida rhombifolia). Dry matter of coffee plants was linearly reduced with increasing B. pilosa and S. rhombifolia density, with pronounced effect of B. pilosa. C. diffusa was the only weed species whose increasing density in the pots did not diminish crop root dry matter. L. sibiricus, N. physaloides and R. brasiliensis reduced root dry matter of coffee plants by 75, 52 and 47%, respectively, as compared to the weed-free treatment, regardless of weed density. Under competition, even though weed species showed lower macronutrient concentration in the roots (except for P), they accumulated 4.2 (N), 12.3 (P), 4.3 (K), 5.5 (Ca), 7.6 (Mg) and 4.4 (S) times more nutrients in the roots than the coffee plants. Crop and weed nutrient concentration, as well as competition degrees greatly varied depending on both weed species and densities.

Weed control under integrated nutrient management systems in faba bean (Vicia faba) production in Egypt

Two field experiments were conducted in two successive seasons, 2005/2006 and 2006/2007, to determine whether management can improve faba bean competitiveness with weeds, thus helping to achieve its yield potential. The experiment included five treatments, composed of organic and mineral fertilizers, alone and mixed at different rates, along with a control and six weed control treatments, including oxadiargyl, prometryn, hand hoeing treatments alone or mixed with the herbicides, and a nonweeded treatment (control).The herbicide treatments were not superior to the two hand-hoeing treatments. Using compost favored growth and yield of faba bean more than of weeds. Adding fertilizer also improved most yield parameters. Application of compost alone or combined with 50 or 100% of the recommended NPK rate improved faba bean growth in terms of net assimilation rate, specific leaf area, and leaf weight ratio as components of relative growth rate. This improvement in growth resulted in increase of seed yield, yield components and protein of faba bean. Faba bean yield performance improved under interactive fertilizer effects and weed control treatments as growth improved, as a result of nutrient release from fertilizers and weed control.

Growth, nodulation, and yield of soybean and associated weeds as affected by weed management

Two field experiments were conducted at the experimental farm of the National Research Centre at Shalakan, Kalubia Governorate, Egypt, during 2006 and 2007 seasons, to study the effect of three pre-emergence herbicides, prometryn [at the rate of 0.75, 1.5 and 2.25 kg ha-1], oxadiargyl [at the rate of 240, 480 and 720 g ha-1] and butralin [at the rate of 1.20, 2.40 and 3.36 kg ha-1], two hand hoeing treatments and a nonweeded check, on weed infestation, nodulation, growth, yield and yield attributes of soybean plants. Two hand hoeing treatments resulted in the highest weed depression expressed as the lowest fresh and dry weights of broadleaved, grassy and total weeds. The reduction percentage in weed dry matter compared to the nonweeded treatment was 98.3, 92.64 and 96.9% in broadleaved, grassy and total weeds, respectively. Application of the three herbicides at higher or recommended doses significantly reduced fresh and dry weight of the weeds compared to the nonweeded treatment. The results indicated that all the three herbicides at rates higher than the recommended markedly decreased the number, fresh and dry weight of nodules as well as root, shoot and total dry weight plant-1, while application of two hand hoeing treatments significantly increased these traits. Two hand hoeing treatments and pre-emergence herbicides at the recommended rates markedly increased soybean yield and its attributes. Two hand hoeing treatments gave the highest values of number of pods per plant-1, weight of pods per plant-1 and number of seeds per plant-1 by 140.7, 150.0 and 59.8%, respectively, compared to the nonweeded treatment. On the other hand, oxadiargyl at the recommended rate (480 g ha-1) was the best treatment for promoting seed yield (g plant-1), seed yield (kg ha-1) and biological yield (g plant-1) compared to the nonweeded treatment by 87.3, 85.0 and 88.2%, respectively. Prometryn at the rate of 1.50 kg ha-1, followed by two hand hoeing treatments, produced the highest shoot and seed protein percentage as well as seed oil percentage, compared to the other weed control treatments.

Weed control via intercropping with gliricidia: I. cotton crop

The majority of cotton grown commercially in the world has white lint, but recently, there has been a growing interest in colored lint cotton in several countries, including Brazil. The use of naturally-colored fiber reduces chemical pollution. The objective of this paper was to evaluate cotton cultivar fiber yield in response to weed control via intercropping with gliricídia. Cultivars BRS-Verde (greenish fibers), BRS-Rubi (reddish brown fibers), BRS-Safira (brown fibers), and BRS-187 8H (white fibers) were submitted to the following treatments: no hoeing, two hoeings (at 20 and 40 days after transplanting), and cotton intercropped with gliricídia. In the intercropped treatment, gliricídia was planted between rows of cotton plants, using one seedling pit-1, in pits spaced 50.0 cm apart. Twelve weed species predominated in the experiment, many of them belonging to the Poaceae family. Weeds occurred at different frequencies and in a non-uniform manner in the experimental area. Cultivars did not influence weed dry matter. Intercropping with gliricídia reduced weed dry matter but did not prevent reductions in cotton fiber and seed cotton yield, which were higher in hoed plots. Cultivar BRS Safira had the highest fiber yield, but no differences were observed between cultivars regarding to seed cotton yield.

Weed control via intercropping with gliricidia: II. corn crop

A reduction in herbicide use is one of modern agriculture's main interests and several alternatives are being investigated with this objective, including intercropping. Gliricídia (Gliricidia sepium) mulch has no allelopathic effect on corn or beans but significantly decreased the population of some weed species. The objective of this study was to evaluate green ear and grain yield in corn cultivars as a response to weed control achieved via intercropping with gliricidia. A completely randomized block design with five replicates and split-plots was used. Cultivars AG 1051, AG 2060, BRS 2020, and PL 6880 (assigned to plots) were submitted to the following treatments: no hoeing, hoeing (performed at 20 and 40 days after sowing the corn), and corn intercropped with gliricidia. Gliricidia was grown in a transplanting system to ensure uniform germination and fast establishment in the field. Seeding was made in 200-cell trays with one seed per cell (35 mL volume). The plants emerged two to three days after sowing and were transplanted to a permanent site two to three days after emergence. Corn was sown on the same day gliricidia was transplanted. Sixteen weed species occurred at different frequencies, with uneven distribution in the experimental area. Cultivars AG 1051 and AG 2060 were the best with reference to most characteristics employed to evaluate green corn yield. Cultivar AG 1051 provided the highest grain yield. The highest green ear yield and grain yield values were obtained with hoeing. However, the fact that intercropped plots showed intermediate yield between the values obtained for hoed and non-hoed plots indicates that gliricidia was beneficial to corn, and exerted a certain level of weed control.

Perspectives for using light quality knowledge as an advanced ecophysiological weed management tool

The current knowledge of light quality effects on plant morphogenesis and development represents a new era of understanding on how plant communities perceive and adjust to available resources. The most important consequences of light quality cues, often mediated by decreasing in red far-red ratios with respect to the spectral composition of incident sunlight radiation, affecting weed-crop interaction are the increased plant height and shoot to root ratio in anticipation of competition by light quantity, water or nutrients. Although the concepts related to light quality have been extensively studied and several basic process of this phenomenon are well known, little applications of photomorphogenic signaling currently are related to agricultural problems or weed management. The objectives of this review are to describe how light quality change can be a triggering factor of interspecific interference responses, to analyze how this phenomenon can be used to predict weed interference, to reevaluate the critical periods of interference concept, and to discuss its potential contribution towards developing more weed competitive crop varieties. Knowledge on light quality responses involved in plant sensing of interspecific competition could be used to identify red/far-red threshold values, indicating when weed control should be started. Light quality alterations by weeds can affect grain crop development mainly in high yielding fields. Unlike the traditional concept or the critical period of competition, light quality mediated interference implies that the critical period for weed control could start before the effects of direct resource (water, nutrients and available light) limitation actually occur. The variability in light quality responses among crop genotypes and the identification of mutants insensitive to light quality effects indicate that this characteristic can be selected or modified to develop cultivars with enhanced interspecific interference ability. Knowledge on light quality-elicited responses represents a new possibility to understand the underlying biology of interspecific interference, and could be used in the development of new weed management technologies.

Genetic variability among Commelina weed species from the states of Paraná and São Paulo, Brazil

This work aims to carry out a comparative analysis using RAPD molecular markers in four Commelina weed species from the state of Paraná and C. benghalensis populations from the states of Paraná and São Paulo, Brazil. The genomic plant DNA sample was extracted from the leaves, separated, randomly fragmented and amplified by PCR. Random amplified polymorphic DNA fragments (RAPD markers) were analyzed by using POPGENE statistical program. Eighty-five primer sequences were tested but only three were suitable as molecular markers producing 37 DNA polymorphic fragments for comparisons among four Commelina species and 22 polymorphic fragments for comparisons among C. benghalensis populations. The results showed that there were inter-specific and intra-specific genetic variabilities among Commelina plant genera. Genetic diversity analysis between species indicated four mono-specific clusters and it was suggested to keep C. villosa as one species. Regarding the intra-specific genetic variability of C. benghalensis alone, three groups were verified, although there were 13 populations from two geographical areas. However, these clusters do not correspond to the distinct characteristics verified.

Effect of cowpea intercropping on weed control and corn yieid

Herbicides have simplified weed control, but the use of herbicides, besides being costly, resulted in the selection of herbicide-resistant weed biotypes and has become an environmental contamination factor. Herbicide use reduction is one of the goals of modern agriculture, with several alternatives being investigated, including intercropping. The objective of this study was to evaluate the effects of cowpea and corn cultivar intercropping on weed control and corn green-ear (immature ears with 80% humidity grains) and grain yield. A completely randomized block design with split-plots and four replications was used. AG 1051, AG 2060 and PL 6880 corn cultivars (assigned to plots) were submitted to the four treatments: no weeding, two hoe-weeding (22 and 41 days after planting), and intercropping with cowpea (BR 14 and IPA 206 cultivars, with indeterminate growth). The cowpea was planted (with corn planting) between the corn rows, in pits 1.0 m apart, with two plants per pit. The corn cultivars did not differ from each other as to weed density (WD), fresh above-ground weed biomass (WB), green-ear yield and grain yields. Higher WD and WB mean values were found in no weeding subplots; lower mean values in two hoe-weeding subplots; and intermediate mean values in intercropped subplots, indicating that cowpea plants had, to a certain extent, control over weeds. The no-weeded plots and the intercropped plots had lower green-ear and grain yields. Although the cowpea cultivars had a certain control over weeds (mean reductions of 22.5 and 18.3%, in terms of green matter density and weight of the above-ground part of weeds, respectively), they also competed against the corn plants, leading to yield reduction (mean reductions of 17.0 and 32% in green ear and grain yield, respectively). The cowpea cultivars did not produce grain, certainly due to the strong competition exerted by the corn and weeds on cowpea plants.