Lambsquarters (Chenopodium spp.) taxa
-Lambsquarters Bibliography (1991)
-Lambsquarters Bibliography (1996)
a. common lambsquarters (Chenopodium album)
b. mapleleaf goosefoot (C. hybridum; C. gigantospermum)
c. mexican tea (C. ambrosoides)
d. taxa: complex of associated species or demography?; Atriplex spp.?
Agronomy 517: Weed Biology and Ecology
Spring Semester, 1996
Kindom Division Subdivision Class Order Family Genus
Chenopodiaceae species: annual or perennial herbs, often fleshy or woody. Leaves:opposite or alternate, entire, toothed or lobed. Flowers:perfect or imperfect and the plant monoecious. Inflorescence: spicate or axillary cymules. Sepals: 3-5 united, persistent, often enlarging in the fruit. Petals: absent. Stames: 5. Fruit: utricle.
Important genera: Chenopodium, Atriplex, Salsola. Provide important species of weeds in USA. Characteristics in filotaxia,leaf shape and fruit make the distintion among genera. About more than 20 species of Chenopodium are distributed in USA and around the world. Several of them considered toxic, for example C. Album cause: a) polen--- summer hay fever; b) contain potentially dangerous concentration of N0-3 which probably in Iowa cause a seasonal disease of swine, perirenal edema to the ingestion of this weed. Another common weed C. Ambrosoides or "wormseed" from tropical America and now with a very wide distribution in North America, it is famous for its antihelminticum properties. The oil responsable for the antihelmintic action (compared by terpene Ascaridol), could be fatal in overdose in animals and people.
Chenopodium genus: Erect, annual or occasionally perennial, monoecious herbs. Leaf alternate, pinnately veined, often 3-nerved,petiolate. Inflorescence of paniculate spikes of cymules, the spikes 0.5-3.0 cm (rarely 10 cm) long. Sepals 5, persistent and tightly enclosing on the fruit. Stamens 5. Stigmas 2-3. Fruit utricule smooth to rugose. Seed horizontal (Radford,et.al.1968).
The most important weed of the entire family and in which this revision will be concentrated.
Geographical origin: Completely obscured as is its relation to other members of its genera for one or another reasons, it does not have readily identifiable centres of diversification. Moquind-Tandon (1840-1849), considered as a weed of agriculture throughout temperate regions.
Geographic distribution: cosmopolitan, ranked amongst the 10 most important weeds on the world Holm, 1977). Occurs in latitudes of 70*N and 50*S.
General description of the species: annual; 10-250 cm. High; stem: angular, branched, brownish-yellow and ridged with green or reddish parallel stripes; leaves: alternate, petioled, without stipules, nearly farinosa to nearly glabrous, deep green to light green, ovate lanceolate to rhombic -lanceolate,sinu-dentate to entire, lamina 1-12 cm.long, 0.5-8.0 wide, 1-5 times longer than wide.Plant: monoecious; flower: perfect, clustered in contiguos glomerules, 5 merous, farinose to glabrous, perianth basally united, clasping and nearly enclosing the mature fruit (utricule); Stames: 5, rarely 3 or 4; pistil having a short style and 2 papillate stigmas; seeds: mostly shiny black, horizontal, 1.2 mm wide by 1.3 mm long, circular in outline; testa with radiating faint rugalate ridges or smooth; pericarp easy or difficult to remove, mottled-farinose or smooth; brown seed, very small proportion; the same size, thin testa regulate ridges or smooth, less heavy.
Account of variation:
Unisexual female and male organs (monoecious) on the same plant.
Wind pollinated --->self and cross pollination.
Genoma ---> Two ploidy levels --> HEXAPLOID 2n=54 most common
TETRAPLOID 2n=36 plus variable number of supernumerary B chromosomes
Atrazine - tolerant biotypes (Canada and USA) = n=27 with completly meiosis.
Aellen,1929; recognized 34 subspecies, varieties and forms in North America, and Abrams in 1944, concluded that all these taxa were minor variants of C. album. In Canada and North America C. Berlandieri 2n=36 and C. Strictum var. Glaucophylus are very related species with C. album
Hybryds: there is not evidence in the current revision that natural interspecific hybrids can occur. Artificially, Darmency and Gasquez obtained Hybrids between the two karyotypes of C. Album. They are mostly sterile and produce seeds of poor viability.
Habitat:--->Climatic growth conditions a) range of annual average precipitation---->30 - 325 cm. b) growing season -->30-200 days. c) range of degree days above 42 *F->1500-3500. Soil conditions --> strongly acid to alkaline, well-drained soils, preferently calcareous soils. Chernozemic, podzol or gleysolic soil type.
Occurrence--->It is found in growing in association with other weeds in disturbed open habitats. It is rare to find it in natural situations such as native woodland and prairie.
Wind-pollinated flowers, probably no insects; self compatible and reproduces both by self and cross-pollination.
Seeds: Seed production capacity >72400/plant and ~ 1428/g
Seed viability in the upper layer (9 in.)of cultivated soil in England = 73 - 12.9 Mill/ac. (Roberst,1968). The extreme phenotipic plasticity make a plant reacts to the environment stress by varying the number of its parts, then an individual plant of C. Album may, under stress of nutrient deficiency; high density or drought conditions, flower and set seed when only it has 5 cm. high, but given more ideal growing conditions, may reach 1.0-1.5 m. hight and produce 50000 times as many seeds as its depauperate stressed counterpart.
Seed dispersion: no obvious mechanism for the dispersion of seed, it has been recorded arising from faeces of animals (cattle, pigs, sheeps, and in birds, seeds pass through their digestive system, increasing germination among 32 to 100%). The size and weight of the seeds allow them to be dispersed efficiently in land-crops by irrigation. Man is the main source of dispersion.
Seed dormancy and seed germination processes:diversefied and complex adaptive mechanims that allow Ch.album to survive and be succesful. It may remain viable in the soil for 1600 years ( Odum,1965 ). It can germinated over a wider range of conditions than the less weedy species of the genus.
Genetic control of innate dormancy: control through "Somatic Polymorphism", common in many Chenopodiaceae species. In Chenopodium album: four types of seed in two colour categories: black and brown and two seed categories: reticulate and smooth. Brown seeds are: thin-walled, larger, germinate quickly even at low temperature. These
are among the first seeds to be ripened by the parent.
Black seeds are: * produced first for the plant under long day lenght; thick-walled, smaller and require cold or supply NO3 to break dormancy (William and Harper,1965).
Seedlings from brown seeds are commonly killed by winter cold or tillage, but, if they survive,produce very large plants with a higher reproductive output than black seeds. The proportion of black (97%) and brown (3%) seed suggest polymorphism is selectively favoured and probably environmental governed as in other genera (Williams and Harper 1965).
Viability of seeds: seeds produced under short days (8h), had 94% and 83% germination in the light and in the dark. Seeds produced under long day (17h), had 20 and 2% germination in the light and in the dark respectively. The inhibitin sustances were different in both photoperiods (8 and 17h.). Germination of 39 years old seeds was obtained by Toole and Brownin 1946, at 45-55 cm., and 90-105 cm. of the soil profile, where conditions of moisture , temperature, and oxygen were assumed to be nearly constant. Lewis,1973 found seed survival buried in soil for 20 years.
Soil Seeds emergence:% accumulated in a 5 years period = 18.5 almost 25 of the total emergenced at the first year (Chepil,1946)
Seeds germination:1- growth regulators - cytochynins and gibberellins in the seed norm this proccess. This aspect was showed by Chu, et. al.,1978.
2- Light requirements: The influence of light on germination is conditioned by many factors and can not be considered as an isolate innate property of the black seeds. The nitrate supply, temperature, age and genotype of the seed are the complex that norms the effect of light on the germination. There are some aspects about, that must be important to be considered:
a) Seed color and dormancy: black seeds- have innate dormancy and need special treatment to germinate. They during the first year harvested have a strong dormancy influenced by such factors as light, moisture, and nitrates and gradually it become reduced as the seed age increase. Nitrate largely determines the response of young seeds to the light, and has more influence than alternating temperature. Constant t* (23*C), ligth and nitrates together promote germination of the young seeds (8-11 months old), which were insensitive to either factors applied separately. Old seeds (32-35 months), show good response to light and nitrate together and separately. After the response to nitrates, lambsquarter has similar features of innately ligth condition like other species: a) short light exposure requirement; b) positive light response with initial water uptake; c) insensitivity of dry seeds; d) decreased light response with prolonged imbibition in the dark. e) light temperature dependence; f) high temperature induction thermodormancy. Brown seeds- present no restriction to germinate in a suitable condition of water, oxygen and temperature (Henson,1970).
b) Spectral composition: the phytochrome system which controls the light sensitivity of germination in many species involves an interconversion of two forms of phytochrome, red light (ca 660nm), and far-red light (ca 730 nm) and probable important in the germination of seeds of Ch. Album could be that far-red light predominates over red under leaf canopies of crops.
d) Interactions: growth regulators and light- in C. album, seeds gradually lose their sensitivity to light and plant growth regulators (hormones) during induction into secundary dormancy. For example, incubating light requiring seeds in darkness led to a gradual decrease in GA4+7 and ethylene. Indicating that there is a gradual transition between non dormancy and dormancy in whatever internal processes light and plant hormones regulate. The germination inhibitors are located in the perianth of fruits that vave been initiated under long photoperiod (20h).
3- Nutrients in the soil.
Nitrogen and related compounds -Fawcet &;Slife, 1978.,found close correlation between rates of N applied and concentration of nitrate in lambsquarters seeds.they found also more seed germination 34%, in plots with previous application of N03NH4 compared
with 3% from unfertilized plots. It may be that stimulation of germination in seeds of C. Album, and a number of other species by nitrate is involved in the timing of field germination, because the NO3 concentration of the soil solution often rises quite sharply as the soil temperature increase in spring.
Oxygen--Lowered oxygen tension has also been held responsible for dormancy of buried seed. Severe starvation may mantain a dormant condition.
Revewing earlier works showed conflicting evidence that just can be explained by the complexity of this process and by the diversity of experimental conditions in which they occurred.
The factors covered in this revision are the most important issues that govern this phenomenon.
Crop-competition effect: the most important weed in potatoes and sugar beet on the world; in United States is very important weed in soybean, asparagus, corn, in Canada, it is important in several crops like: soybean, cereals (wheat, oat, barley etc.); european countries, England, French, germany, etc., mainly in cereal crops; in Mexico, commonly found in cereal crops, corn, beans and in the establishment of alfalfa.
1- Recruitment of seedling population: High density of seedlings are normally found in crop fields early in the spring ( 0.8 - 0.2 mill/ha.), and they can continued germinating until late of the summer. As soon the crop or weed canopy start to cover or shaded the soil, the germination is inhibited. Microtopography had a marked effect on the relative establishment of the species, the behavior on the soil surface could be seen in the way in which it comes to the rest of species after sowing. The experiments established at this respect, emphasize a somewhat unusual aspect of seed size and shape as well as a neglected element of environmental variation in determining the size and quality of plant population.
3- Density-Mortality relationship: When plant population are grown at high density, some individuals often die. There are two categories according with the type of relation established among individuals in a population: a) density dependent or self-thinning: the mortality of some individuals in a population is due to the pressure of individuals of the same species, and b) density independent or alien -thinning: where the mortality of individuals from a specific species can be ascribed to the streess from the density of an associated species. C. Album shows a self-thinning relationship, where the survivors plant presented a negative relationship between the weight of survivors and density.
2- Weed -to- weed competition:
Redroot pigweed (Amaranthus retroflexus L.) and Common Lambsquarters ( Chenopodium album L.) are common associates in the weed community that grow in many parts of the agriculture's world. Their behavior in the field,growing together, can be expected to vary from year to year,mainly according with the temperature in the early period of their association, and this temperature is related by the time of planting. Interesting work was done by Chu, et. al.1978b, finding:
Pigweed germinates: much more rapidly than lambsquarters under higher temperature.
much more slowly than lambsquarters under the lowertemperature.
Lambsquarters grow: the same like pigweed under lower temperature
slower than pigweed under higher levels.
If either weed has a time advantage over the other, it completely suppresses the later planted species.
In midwest dry spring,early planting, cool temperature ---Lambsquarters problem
wet weather, late planting, warmer temperature---Pigweed problem
This confirmed that C3 plant C. album was better adapted to shade and cool temperature in fertilized fields than A. retroflexus, a C4 species. Corn a C4 plant suffers severe competition by early germinating C. album under the climatic conditions of central Europe. The C4 weeds A. Retroflexus and E. Crusgalli, only have chance if Chenopodium is removed. In this case , time of germination is most crucial for success in comprtition and not the C3 orC4 pathway.
The differences in germination and growth of the two species in response to temperature, explain their competitive ability,remember us that data on growing conditions are needed for valid comparison of weed population among experiments (Aldrich, 1984). The makeup of the weed community to change somewhat from year to year is a common situation even under monoculture and even though the seed bank become constant from year to year, this just could be explained by the way in which each member of the community sense the environment in a specific term of time and space. The relationship between lambsquarters and redroot pigweed is a very didactic example of the what was said before.
Response to weed control practices
Extensive inputs of tillage and herbicides are used to control C. album, mainly in summer annual crop production system around the world. The crop production system environment could be much of times characterized by high fertility, annual disturbance, production in rows, and herbicide use. In these scenary of conditions, the dynamic of population of C. album, respons in a successful or failed form, according with its biological characteristics interacting with the efficacy of weed management practices.
Polymorphism as a response to the tillage practices
The general responses of common lambsquarters to the tillage presure is inconsistent and some times contradictory. The classic studies of Roberts and his team, showed that, if all seeding of weeds is prevented, then progressive reduction in the number of viable seeds in the soil accurred due at tillage effect. The rate at which it occurs will depend of the species, frequency, and time of cultivation. In C. Album, it was 40% and it was less affected for more than one culivation. They also found that the viability of the Lambsquarters seeds decresed more in a Not-Tillage condition. Results showed by Buhler (1995),indicated that in soybean grown in rotation with corn, common lambsquarters densities were affected by tillage at one location, but not at another location with similar edaphic and cropping history, where no tillage land showed less density. Probably the explanation given by Fawcet (1987) could help in understanding this facts. He said that species that germinate under cool soil conditions, may be prevalent in no tillage fields prior to crop planting. This may allow them to suppress later emerging weeds or a significant portion of the population may be destroyed by tillage or herbicides prior to crop planting. Therefore, the effects of planting date and spring soil temperature in C. Album, germination prior to crop planting , probably caused the inconsistent responses observed among studies (Buhler,1995).
Polymorphism and herbicide resistant
Resulted of the intensive use of herbicide in the C. Album population, soon brought the appearance of biotypes resistant to the triazine group of herbicides.
The repeated use of triazine herbicides assumably has favored the selection of existing resistant plants or mutants. Darmency and Gasquez (1990)show out a revision of appearance and spread of triazine in C. album, indicating that, the more polymorohic the population is, the more likely that it can adapt to new agricultural stress.
One aspect of the polymorphism of this species,is the simultaneous presence of two ploidy levels in each population.the most common karyotype is hexaploid, 2n=54, corresponding to the species voucher type; the other is tetraploid, 2n=36 plus a variable number of supernumerary B chromosomes. The tetraploid type does not correspond to any described species in Europe. As it can not be distinguished from typical C. album by its morphology but flower sooner, it can not correspond to the tetraploid lateflowering goosefootC. Strictum Roth. var. glaucophyllum (Allen) H.A. Wahl. For which Atrazine-resistant population were found in Canada.
However, the two ploidy levels belong to the same biologycal species, they do not interchange genes in the field because the carrying pollen by the wind is very resticted in this plants. Thus, low heterozygosity may be expected so that a field population can be considered as just a mixture of different genoypes evolving separately. Due to the low outcrossing rate in corn fields, no gene exchange has occurred between resistant plants and adjacent susceptible population.
Another side of the polymorphism of C. Album populations is revealed by isozymes analysis. Using no more than four enzymes, nearly 60 different types of hexaploid plants in a survey of just two population in France, with an average of 23 genotypes per population.
The appearence of resistant plants within population of C. album, is not at random but depends on the presence of some unusual genotypes that can give rise to psbA gene- mutants with a higher frequency than expected with random mutation alone. It shows one of the most widespread resistant infestation.
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