Seed germination
Common Lambsquarters (Chenopodium album) Seed Heteroblasy:
Morphology, Environment and Germination
Two publications provide insight into the germination structure of common lambsquarters seed rain arising from seed morphology and environment, with implications for the patterns of seedling recruitment.
Seed polymorphism and germination (Williams and Harper, 1965)
The effects of light, temperature, after-ripening, nitrate and water on Chenopodium
album seed germination (Altenhofen, 2009)
The effects of light, temperature, after-ripening,
nitrate and water
on Chenopodium
album seed germination (Altenhofen, 2009)
Seed
heteromorphy
Chenopodium album flowers
are arranged in glomerules, forming spikes at a terminal panicle (Gleason and Cronquist,
1963). Seeds appear to be disk-shaped with two
convex sides. Detailed seed morphology and classification is not agreed upon in published
literature, but similarities are evident among descriptions as illustrated herein. As described by Delorit (1970), seeds of C. album
are black, with a glossy surface and enclosed in a semi-translucent granular pericarp.
The five-parted perianth may also be attached to the seed, but is often detached during
collection and subsequent handling. The seeds of C. album have been described by
Williams and Harper (1965) as having four distinct forms at maturity: brown-reticulate,
brown-smooth, black-reticulate and black-smooth. Additionally,
all of these seed types may be present on the same plant. Chu, Sweet and Ozbun (1978)
described seed as having four color morphs: brown, deep brown, brown-black and black but
they did not distinguish between smooth or reticulate. Wentland (1965) described the
variation in color as yellow, orange, and dark red being characteristic of immature seed,
whereas mature seeds are dark in color. A few studies considered here (Henson, 1970;
Moravcova and Dostalek, 1989) stated that they used only black seed while most studies did
not specify seed morphology.
The study reported
herein used only seed which appeared to be brown to dark brown in color, with pericarp
intact or nearly complete, as this was the state of seed when collected. A perianth was
present on roughly half of seeds used, in an open or closed formation, as collected in the
field. Immature, yellow, orange or red seeds were omitted from experiments.
Morphology and germination
Germination tests were
conducted in the light on the several morphological types of C. album seed
present in Ames, Iowa populations (October 5, 2007 harvest; Table 1). The structure of C. album seed variability, seed heteroblasty, was
revealed in the differences among seed morphological phenotypes to alternating
temperature.
Table 1. Structure of Chenopodium album seed germination (%) in
response to seed hull morphology (color, structure, maturity), alternating temperature (5-15; 15-25; 25-35° C), and chilling pretreatment
(4° C-wet-dark after-ripening; 0-35 days).
Seed Hull Phenotype |
Alternating Temperature |
Maximum % Germination |
|
Color |
Morphology |
||
Red |
immature |
5-15° C |
15 |
15-25° C |
5 |
||
25-35° C |
20 |
||
Dark |
sepals and perianth present |
5-15° C |
5 |
15-25° C |
20 |
||
25-35° C |
10 |
||
reticulate (net-like) hull |
5-15° C |
10 |
|
15-25° C |
15 |
||
25-35° C |
5 |
||
smooth hull |
5-15° C |
35 |
|
15-25° C |
45 |
||
25-35° C |
15 |
||
The greatest germination was observed with dark-smooth
seeds, which also were the most responsive to cool (5-15° C) and warm (15-25° C)
temperatures. Immature red seed were the most
readily germinable (data not presented) and most responsive to hot (25-35° C)
temperatures. Among dark seeds, germination
was less with reticulate relative to smooth seeds.
Environment and germination
The structure of C.
album seed variability, seed heteroblasty, was revealed by conducting germination
assays on propagules collected in Ames, Iowa in 2007 (October 5) and 2008 (October 11). These seed germination assays were organized as a
factorial arrangement of treatments to understand the complex interactions of 6
parameters:
1] Population
(year of collection),
2] Chilling
after-ripening (4° C-wet-dark) pretreatment (0-10 weeks) prior to the germination assay,
3] Light
regime (light or dark),
4] Alternating
diurnal temperature (5-15; 15-25; 25-35° C),
5] Nitrate
(+ or -),
6] Water
quantity (0.75, 1.0, or 1.25 ml)
All six parameters affected C. album seed germination, but groups of seeds
within each population responded to these stimulatory factors differently (Table 2). The germination structure, seed heteroblasty, was
made apparent in different combinations of these factors.
There exist different germination fractions in C. album seed populations: the seed rain consists of different groups, each of
which is stimulated by different sets of germination conditions. Germination was stimulated primarily by light, but
a small fraction was revealed in dark germination. Germination
was greatest in warm (15-25° C) alternating temperatures, but other fractions responded
to hot (25-35° C) conditions. Nitrate
stimulated germination significantly, but some fractions responded well to just water. In fact the 2008 population germination increased
with increasing water quantity (warm; nitrate), a fractional response not observed in the
2007 population. Fine scale germination
structure was revealed in many factor combinations with increasing dosages of pre-chilling
(oxygenation): each week (0-4 or 5) more germination sub-fractions were revealed.
Table 2. Structure of Chenopodium album seed germination in response to
year of embryogenesis, photoperiod (light, dark), alternating temperature (5-15; 15-25; 25-35° C), nitrate (0.01 M),
water quantity (0.75, 1.0, or 1.25 ml), 4° C-wet-dark after-ripening (chilling) duration
(weeks) and percent seed germination; -, averaged over this parameter of the
interaction.
Year |
Light-Dark |
Alternating Temp |
Nitrate (0.01 M) |
Water Amount |
Chilling Time to Max Germ |
% Seed Germ |
2007 |
Dark |
- |
-NO2 |
- |
- |
0-2 |
Light |
- |
H2O |
0-4 weeks |
2-7 |
||
-NO2 |
- |
7-22 |
||||
25-35° C |
- |
0-5 weeks |
5-20 |
|||
15-25° C |
5-25 |
|||||
2008 |
Dark |
- |
- |
- |
- |
0-2 |
Light |
25-35° C |
H2O |
= |
0-4 |
||
15-25° C |
0-4 weeks |
4-19 |
||||
25-35° C |
-NO2 |
- |
3-38 |
|||
15-25° C |
0-5 weeks |
6-50 |
||||
0.75 ml |
- |
27 |
||||
1.00 ml |
33 |
|||||
1.25 ml |
38 |
|||||
Conclusion
The complexity of C. album seed germination structure may provide
some insight into the complexity of recruitment patterns revealed in the research
conducted by the European Weed Research Society, Working group on germination and early
growth, on this species.
References
Altenhofen, L.A.
2009. The effects of light, temperature, after-ripening, nitrate and water on Chenopodium album seed germination. M.S. Thesis;
Ecology and Evolutionary Biology, Iowa State
University.
Chu, C. C., R. D. Sweet, and J. L. Ozbun. 1978. Some
germination characteristics in common lambsquarters (Chenopodium album). Weed Sci.
26:255-258.
Delorit, R. J. 1970. An illustrated taxonomy manual of
weed seeds. Agronomy Publications, River Falls, Wisconsin. p 76.
Gleason, H. A. and A. Cronquist. 1963. Manual of
vascular plants of northeastern United States and adjacent Canada. D. Van Nostrand
Company, New York, New York. p 274.
Henson, I. E. 1970. The effects of light, potassium
nitrate, and temperature on the germination of Chenopodium album L. Weed Res.
10:27-39.
Moravcova, L. and J. Dostalek. 1989. Contribution to the
biology of germination of four species of Chenopodium album agg. under different
conditions. Folia Geobot. Phytotx. 24:431-439.