More info for the terms: bulb, bulbil, culm, frequency, perfect, serpentine soils
Although bulbous bluegrass produces true florets as well as bulbils, seeds are rare [30,47,80]. Bulbous bluegrass regeneration is almost exclusively through asexual means including bulbil production [10,47], basal bulb production [47], and sometimes through culm rooting [12].
Reproduction through true floret and seed production does occur in bulbous bluegrass's native habitats. Some bulbous bluegrass populations reproduce primarily by seed, while in others reproduction is dominated by basal bulb growth and/or bulbil production. Researchers found a significant negative linear relationship (r=-0.596, P=0.02) between percentage of flowering plants and mean annual precipitation of habitats when plants were taken from field sites and grown in controlled conditions [59].
Pollination: When true florets are produced, they are wind pollinated [60].
Breeding system: Spikelets are perfect [53]. Although asexual regeneration is predominant, researchers found "surprisingly high levels of genetic diversity at the species and population level" when bulbous bluegrass bulbils were collected and grown from 10 populations in Idaho, Oregon, and Washington. An average of 26 plants were grown from each population. Researchers concluded that bulbous bluegrass was an autopolyploid and that the high levels of diversity resulted from multiple introductions and/or some past sexual reproduction [58].
Seed production: Through controlled experiments on bulbous bluegrass strains from Turkey and Afghanistan, Youngner [88] determined that floret and bulbil production were controlled by day length and temperature. Plants grown under high temperature (70-81 °F (21-27 °C)) and long day (16 hours of light) conditions produced normal florets. Plants produced bulbils when grown at lower temperatures (68 °F (<20 °C)) and exposed to shorter days (8 hours of light). Plants exposed to long day and low temperature or short day and high temperature conditions produced panicles with both florets and bulbils. The researcher recalled observations of bulbous bluegrass plants near Los Angeles, California, producing normal florets in late March, when day length exceeded 13 hours [88].
Seed dispersal: Animals, wind, and gravity are all possible dispersers of bulbous bluegrass. Small mammals cache and disperse bulbous bluegrass bulbils [24]. Because leaves and roots senesce with the dry season, it is possible that basal bulbs may be moved by wind (Sowerby cited in [26]). However, this method of dispersal is not described elsewhere, and Halperin [26] indicates he never observed this type of dispersal.
Roadsides are important dispersal corridors for bulbous bluegrass. The frequency of bulbous bluegrass was compared on sites with serpentine and nonsepentine soils at 30, 300, 3,000 feet (10 m, 100 m, and >1,000 m) from paved or improved roads. Bulbous bluegrass did not occur on serpentine soils but had 9.6% and 9.2% frequency on nonserpentine soil sites 30 feet (10 m) and 300 feet (100 m) from roads, respectively. Bulbous bluegrass was not found 3,000 feet (1,000 m) away from roads [20].
Seed banking: Studies regarding the longevity of bulbous bluegrass bulbs or bulbils are lacking. Bulbils attached to the plant that make contact with the soil can root and grow immediately without a dormancy period. Bulbils detached from the panicle lie dormant for 3 to 6 months before germinating [88]. Basal bulbs may live dormant for "several" years [24]. Basal bulbs can dry to below 10% moisture yet maintain growth potential for "extended periods" [83]. Depth of burial affects bulbous bluegrass emergence; for more on this topic, see the Depth of burial discussion.
Germination: Several studies of bulbous bluegrass germination under controlled conditions suggest that germination is best when temperatures are cool 41 to 64 °F (5-18 °C) for at least part of the day [36,58] and when seeds are uncovered or only shallowly covered with soil [63,81]. Bulbous bluegrass bulbils collected from populations in Oregon, Washington, and Idaho usually germinated within 24 hours after being chilled for 4 to 5 days at 41 °F (5 °C) [58]. Germination was 98% to 99% for bulbils collected from Moscow, Idaho, exposed to cool (50 to 64 °F (10-18 °C)) 8-hour days and cold (38 °F (3.5 °C)) nights. No bulbils germinated when exposed to 14-hour warm (79 °F (26 °C)) days and warm nights (68 °F (20 °C)). When day length was shortened to 11 hours under the warm day and night treatments germination was 18% [36].
Bulbous bluegrass bulbils from Intermountain seed suppliers germinated best under fluctuating greenhouse temperatures that ranged from 50 to 100 °F (10-40 °C), averaged 57 °F (14 °C), and closely approximated outdoor early spring conditions. Fluctuating greenhouse temperatures produced 98% germination in 14 days. Room temperature conditions that averaged 70 °F (21°C) and ranged from 66 to 73 °F (19-23 °C) produced 91% germination after 19 days. Germination was only 1% after 24 days at a constant temperature of 86 °F (30 °C) [63].
Depth of burial: Shoot emergence was delayed and reduced as bulbous bluegrass bulbil planting depths increased. Bulbils were kept moist in a greenhouse where temperatures averaged 64 °F (18 °C) and ranged daily from 54 to 90 °F (12-32 °C). Just 8% of seedlings emerged after 30 days when bulbils were planted 1.5 inches (3.8 cm) deep. Emergence was more rapid and more successful when seeds were left of the soil surface or shallowly planted. Researchers indicated that field conditions may not provide adequate moisture for successful germination of uncovered bulbils. A summary of the findings is provided below [63].
Percentage shoot emergence 5 to 30 days after planting at depths of 0 to 1.5 inches [63] Days after planting Planting depth (inches) 5 7 9 11 13 15 30 surface 26 64 85 90 93 95 96 0.25 0 6 47 71 83 87 90 0.50 0 2 26 60 68 73 77 0.75 0 0 15 49 58 62 64 1.0 0 0 0 5 15 22 28 1.5 0 0 0 1 2 5 8
Seedling establishment/growth: Bulbous bluegrass plants produced the most growth in late April and May and senesced by 6 June in a mountain brush field nursery site in the Intermountain Region. The nursery occurred at 5,375 feet (1,638 m) on deep clay loam soils. Bulbils were sown in late fall, and root and shoot development was monitored periodically from 22 March to 6 June. Root and shoot growth was underway in early April. By mid-May there were 70 roots that penetrated 4.7 inches (12 cm) and spread 3.5 inches (9 cm), and shoot height reached 18 inches (46 cm). By 6 June, the approximate start of summer drought conditions, bulbous bluegrass was senescing [63].
Bulbous bluegrass root and shoot growth over the growing season [63]
Sampling date
April 5 April 12 April 18 April 25 May 2 May 9 June 6 Root length (cm) 10 10 14 30 41 73 9 Shoot length (cm) 5 6 10 16 21 46 0 Number of roots 9 9 28 35 37 70 6 Number of leaves 2 3 3 7 9 18 0 Root penetration (cm) 4 4 5 6 8 12 0 Root spread (cm) 2 2 3 7 6 9 0
Vegetative regeneration: Asexual or vegetative regeneration is the primary means of reproduction in US bulbous bluegrass populations. Bulbils produced in the panicle through the transformation of flowers and lemmas germinate to produce new individuals [10,22,47]. Basal bulbs that multiply and grow laterally lead to vegetative spread [47,64]. Vegetative regeneration is also possible through the rooting of decumbent culms [12].