Response of Bluebunch Wheatgrass and Medusahead to Defoliation

Our objective was to determine the short-term response of bluebunch wheatgrass and medusahead to defoliation of wheatgrass designed to stimulate regrowth through tillering. We hypothesized that defoliating bluebunch wheatgrass by 20% at the 3 to 3.5 leaf stage followed by a 50% defoliation at peak standing crop would increase its tillering and biomass production. Consequently, we expected a reduction of the density and biomass of medusahead over that of bluebunch wheatgrass defoliated 50% at peak standing crop. Treatments included four initial medusahead densities (200, 333, 444, 600 plants · m^-2) created by hand-pulling and three defoliation regimes factorially arranged (12 treatment combinations) in a randomized complete-block design and replicated four times at two sites. In 2006 and 2007, defoliation was accomplished by hand-clipping bluebunch wheatgrass 1) by 50% once at peak standing crop (late June); 2) by 20% at the 3 to 3.5 leaf stage, then again to 50% at peak standing crop (mid May, late June); or 3) plants were not clipped. Density was sampled in 2006 and 2007, and biomass was harvested only at Star Mountain (near Riverside, Oregon) in 2007 because Warm Springs (near Drewsey, Oregon) was burned by a wildfire before final 2007 data could be collected. In 2006, no treatments applied at either site detectably altered the number of tillers produced by bluebunch wheatgrass nor did they affect bluebunch wheatgrass density or biomass in 2007 at Star Mountain. Changes in medusahead density were not detected in 2006, but this annual invasive grass increased in density and biomass in 2007 at Star Mountain in plots receiving two defoliations. The relatively short growing period caused by summer drought and the relative intolerance of bluebunch wheatgrass to grazing make the twice-over grazing an unlikely practice for arid rangelands in the western United States. In fact, it could possibly increase the risk of annual grass invasion.     

Above-Ground Net Primary Production for Elymus lanceolatus and Hesperostipa curtiseta After a Single Defoliation Event

Above-ground net primary production (ANPP) of northern wheatgrass (Elymus lanceolatus [Scribn. & J. G. Sm.] Gould) and western porcupine grass (Hesperostipa curtiseta [Hitchc.] Barkworth) was determined after defoliation to a 7.5 cm stubble height on five landform elements in the Northern Mixed Prairie that had been ungrazed for > 25 yr. Landform elements included north aspect–concave slopes, north aspect–convex slopes, south aspect–concave slopes, south aspect–convex slopes, and level uplands. ANPP was determined for 2 yr after defoliating plots once in May, June, July, August, September, October, November, or April. Northern wheatgrass and western porcupine grass ANPP varied among landform elements (P < 0.01), but not with the month of defoliation × landform element interaction (P ≥ 0.22). Month of defoliation did not influence ANPP of northern wheatgrass (P ≥ 0.69), but that of western porcupine grass was reduced by August and September defoliations (P < 0.01). ANPP of both grasses was insensitive to landform element in terms of defoliation responses. Northern wheatgrass ANPP was not responsive to temporal aspects of a single defoliation. With the exception of August and September defoliations, western porcupine grass also was insensitive to a single defoliation in different months. Land managers should consider rest (1 yr nongrazing) following grazing of western porcupine grass in August or September, whereas responses to defoliation in different months suggest northern wheatgrass can be grazed annually.     

Biomass Production and Net Ecosystem Exchange Following Defoliation in a Wet Sedge Community

Riparian ecosystems provide many ecosystem services, including serving as an important forage resource for livestock grazing operations. We evaluated defoliation impacts on above- and belowground production, and net ecosystem exchange of CO₂ (NEE), in a wet sedge (Carex nebrascensis Dewey)-dominated plant community. In June or July of 2004–2005, experimental plots were clipped to 10 cm stubble height and paired control plots left unclipped. All plots were clipped to 2.5 cm in mid-September, and end-of-season and season-long aboveground production calculated. Root ingrowth cores were used to estimate annual root production and root length density (RLD). A portable gas exchange system and plexiglass chamber were used to measure NEE in 2005. An elevated water table in 2005 vs. 2004 was associated with higher (P &spilt; 0.001) season-long aboveground production (about double), but lower (P ≤ 0.05) belowground production (about half). Total productivity did not differ between years, but below-:aboveground ratios were 3× higher in 2004 vs. 2005. RLD was not different between years (P &spigt; 0.05). Clipping reduced (P ≤ 0.05) end-of-season aboveground standing crop by 33% to 73% depending on clipping month and year. Effects of clipping month on season-long aboveground production were inconsistent between years; June clipping decreased (P ≤ 0.05) production (-10%) in 2005 and July clipping decreased (P ≤ 0.05) production (-25%) in 2004. NEE for June-clipped plots recovered within 1 mo of clipping, whereas NEE for plots clipped in July remained below unclipped levels at the end of the growing season. Water table levels strongly influenced below-:aboveground ratios, although total production was relatively stable between years. Year effects overwhelmed clipping effects on season-long aboveground production. Defoliation after mid-summer did not allow recovery of photosynthetic capacity by the end of the growing season, suggesting the potential for long-term impact with regular late-season defoliation.     

Crested Wheatgrass Defoliation Intensity and Season on Medusahead Invasion

The objective of this study was to determine the effects of crested wheatgrass (Agropyron cristatum [L.] Gaertn.) defoliation intensity and timing on medusahead density and biomass. We hypothesized that crested wheatgrass defoliation greater than 60% during the spring would provide maximum medusahead (Taeniatherum caput-medsae [L.] Nevski subsp. asperum [Simk.] Melderis; taxonomy from US Department of Agriculture) density and biomass. Eighteen treatments (six defoliation levels, three seasons of defoliation) were applied to 2-m² plots in a randomized complete block design on two sites with varying clay content. Blocks were replicated five times at each site. Plants were clipped in 2004 and 2005. Crested wheatgrass was hand clipped to defoliation levels of 0%, 20%, 40%, 60%, 80%, and 100% in the spring, summer, or fall. Density of crested wheatgrass and medusahead was sampled in June 2005 and 2006, but their biomass was harvested only in 2006. Data were analyzed with least square means analysis of variance. Over the two seasons, site had much more of an impact on medusahead invasion than either defoliation intensity or timing of defoliation. The results support previous suggestions that clayey soils favor medusahead and that perennial grasses with high biomass can resist this invasive species. On the clayey site where medusahead did persist, fall defoliation of crested wheatgrass reduced the density of this invasive species by 50% or more compared to spring defoliation. Given the developmental pattern of medusahead, the goal of any management program should be to maximize resource use by the desirable species during April to late July.     

Defoliation Timing Effects on Spotted Knapweed Seed Production and Viability

Spotted knapweed (Centaurea stoebe L.), a perennial invasive forb that reproduces largely by seed, often forms new flowers after prescribed sheep grazing or mowing is applied during the bolting or flowering stage. It is unknown if these new flowers produce viable seeds by the end of the growing season. The purpose of this 2-yr study was to determine the appropriate timing (or timings) or combination (or combinations) of timings of defoliation on spotted knapweed to reduce its viable seed production. Spotted knapweed plants on foothill rangeland in west-central Montana were hand-clipped at seven different timings and frequencies of defoliation: June (bolting stage); July (late-bud–early flowering stage); August (full-flowering stage); June + July; June + August; July + August; or June + July + August. Unclipped plants were controls. Plants clipped in the bolting stage were defoliated at 35–40% relative utilization. Plants clipped at all other timings had 100% of their buds and flowers removed, plus 3 cm of each bud or flower stem. Plant response was evaluated from mid-August through September, whenever the seed heads of each treatment’s plants reached maturity but while their seed-head bracts remained tightly closed. Clipping at any timing or combination of timings reduced the number of buds and flower heads per plant (P < 0.01), number of seeds per plant (P < 0.01), percentage of viability of seeds (P < 0.01), and number of viable seeds per plant (P < 0.01) compared with no clipping. Clipping during the bolting stage reduced the number of viable seeds by nearly 90% compared with no clipping. Clipping during the late-bud–early-flower or full-flower stage reduced the number of viable seeds by nearly 100% compared with no clipping. Spotted knapweed defoliation via prescribed sheep grazing or mowing in summer should suppress viable seed production of spotted knapweed.     

Yield Response of Needle-and-Thread and Threadleaf Sedge to Moisture Regime and Spring and Fall Defoliation

Little information is available to help managers of cool-season dominated semiarid rangelands determine when to begin and end grazing in the spring and fall. Therefore, we evaluated the effects of clipping spring and fall growth on subsequent-year yield of needle-and-thread (Hesperostipa comata [Trin. & Rupr.] Barkworth) and threadleaf sedge (Carex filifolia Nutt.) (USDA-NRCS 2012) using a randomized complete block, split-plot experimental design with fall moisture regimes (ambient or supplemental water) applied to main plots and defoliation treatments applied to subplots. Two combinations of spring defoliation, one for each fall moisture regime, were composed of a factorial array of three spring clipping dates (early May, late May, mid-June) and three levels of defoliation (0%, 40%, 80%). A third combination of treatments was composed of the supplemental water regime and an array of a single spring clipping date (late May), a single fall clipping date (late September, after regrowth), and three levels of defoliation (0%, 40%, 80%) in the same year. Ambient fall moisture was low, leading to continued senescence of needle-and-thread and threadleaf sedge, whereas the application of 10 cm of supplemental water in mid-August stimulated fall growth. The study was replicated with two sets of main plots at four sites in consecutive years, 2002 and 2003. Yield data were collected in mid-June of the year following treatment. Subsequent-year yield of needle-and-thread was not affected by defoliation under average plant-year precipitation conditions (2003) (P > 0.05); however, it was reduced following heavy (80%) late spring (late May or June) defoliation during a drought year (2002) (P > 0.05). Subsequent-year yield of threadleaf sedge was not affected by defoliation in either year (P > 0.05). Because it is difficult to predict when drought will occur, avoiding heavy late-spring grazing in needle-and-thread–dominated pastures in consecutive years would be prudent.     

紫花苜蓿刈割和晾晒技术研究

紫花苜蓿（Medicago sativa L.）是我国种植面积最大的豆科牧草,但在常规的收获制度中因收获技术不当往往造成其品质下降和产量降低。为研究不同刈割留茬高度和不同晾晒时间对苜蓿干草品质的影响,将苜蓿草按留茬高度为0,2~5,5~8,8~11 cm进行刈割,并对其草产量、茎叶比、越冬率、水分散失规律及营养价值进行测定分析。结果表明：刈割留茬高度为2~5 cm时不仅能获得高产的苜蓿草,且其后2茬的草产量也显著高于其他留茬高度的（P<0.05）,具有较高的持续生产能力;同时其粗蛋白（CP）含量和相对饲喂价值（RFV）较高。末茬苜蓿草的刈割留茬高度应在8~11 cm,此条件下其翌年的返青率（77.44%）显著高于其他留茬高度下的（P<0.05）。苜蓿经72 h晾晒至安全含水量时,叶片脱落率达56%,而在含水量降至52%时叶片损失率仅为29%,即后期出现严重茎叶干燥不同步现象。基于此,建议苜蓿干燥过程中采用茎秆压扁技术。     

Freezing Stress Influences Emergence of Germinated Perennial Grass Seeds

In sagebrush rangelands perennial bunchgrasses are typically seeded in fall and a high proportion of planted seeds germinate prior to winter onset but fail to emerge in spring. Our objectives were to evaluate freezing tolerance of germinated but nonemergent bluebunch wheatgrass seeds under laboratory conditions. We used data from a 2-yr pilot study to determine overwinter freezing temperature and duration for soils in southeastern Oregon. We then conducted two experiments to assess freezing tolerance. In experiment 1, bluebunch wheatgrass seeds were planted in control pots and compared to seeds planted at early, mid, or late postgermination stages. Pots from each treatment were placed in a grow room maintained at 12 h 40 min light/11 h 20 min dark photoperiod, with a constant temperature of 22°C for 30 d either immediately or following a 30-d freeze. In experiment 2, germinated bluebunch wheatgrass seeds were planted in pots that were left nonfrozen or were frozen for a specified duration prior to a 30-d period in the grow room. Emergence density and tillers · seedling^?1 were quantified for both experiments. The number of days per year for freezing soil conditions in the pilot study ranged yearly from 25 to 51; maximum duration of continuous freezing was 16.5 and 11.2 d. Freezing reduced or eliminated seedling emergence at all postgermination stages (P < 0.001) and tiller density was reduced by at least 50% (P < 0.001). Maximum reduction in seedling density (P < 0.001) was realized within 4 d of initiation of freezing and tillers · seedling^?1 were reduced 30–70% with > 6 d of freezing (P = 0.001). Our data indicate that freezing-associated mortality of germinated but nonemergent bluebunch wheatgrass seedlings can be extremely high and suggest that management practices to reduce prewinter germination of seeds could improve subsequent emergence and seeding success.     

Plant Establishment in Masticated Utah Juniper Woodlands

Juniper (Juniperus spp.) encroachment into sagebrush (Artemisia spp.)-bunchgrass communities has reduced understory cover on millions of hectares of semiarid rangelands. Mechanical masticators shred trees to restore desirable vegetation and reduce the potential for catastrophic wildfire. Mechanical mastication where juniper density is high and perennial grass cover is low brings a risk of invasive weed dominance unless perennial species are established. To determine whether juniper mastication favors annual- or perennial-grass establishment, we compared seedling emergence, tillers, and aboveground biomass of cheatgrass (Bromus tectorum L.) and Anatone bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Löve). Comparisons were made among hand-planted rows between and under juniper canopies of masticated and adjacent untreated control areas at three locations in Utah. Bluebunch wheatgrass had 16% (95% CI: 11–21) and cheatgrass had 10% (95% CI: 5–15) fewer seedlings emerge per row in masticated than untreated areas (P < 0.001). However, bluebunch wheatgrass had 3.2 (95% CI: 2.0–5.2) times more tillers and 1.9 (95% CI: 1.6–2.2) times more aboveground biomass per row in masticated than untreated areas (P < 0.001). Similarly, cheatgrass had 2.3 (95% CI: 1.5–3.8) times more tillers, 2.0 (95% CI: 1.7–2.4) times more aboveground biomass, and 11.4 (95% CI: 6.3–20.7) times more spikelets per row in masticated than untreated areas (P < 0.001). This increased seedling growth in masticated areas was associated with increased inorganic nitrogen and soil water compared to untreated areas. Because mastication improves the growth of both cheatgrass and bluebunch wheatgrass seedlings, it could support dominance by either annual- or perennial-life forms. To avoid cheatgrass dominance where perennial understory cover is limited and cheatgrass propagule pressure is high, mastication should be accompanied by seeding desirable perennial species such as Anatone bluebunch wheatgrass.     

刈割对热研13号柱花草光合特性的影响

在盆栽条件下,用20,40和60d3个刈割周期与5,15和25cm3种留茬高度组合为9个刈割处理,对热研13号柱花草的光合特性进行研究。结果表明：刈割周期和留茬高度的互作对柱花草光合特性的影响,气孔导度（Gs）和蒸腾速率（Tr）达到显著水平,其他指标均不显著。高频次低留茬（20d5cm）刈割处理的柱花草光合速度（Pn）,Gs,胞间C02间浓度（Ci）和Tr均最高,分别达到了20．52umol CO2／（m^2·s）,0．49umol／（m^2·s）,318．89uL／L和11．32mmol／（m^2·s）。而低频次高留茬（60d25cm）刈割处理柱花草的WUE最高,达到了2．75umol CO2／（mmol·H2O）。     

刈割高度对一年生牧草再生性能及产量的影响

通过对哈尔滨市几种一年生牧草不同刈割高度下,再生性能及产量的研究,结果表明,刈割时留茬高度不同,对牧草的产量有显著影响。晚熟籽粒苋留茬50 cm,刈割时期为抽穗初期,刈割次数可达到3次,总产量可达到54 500 kg/hm^2;晚熟苦荬菜株高达到60-100 cm,留茬20 cm,产量达到43 950kg/hm^2;菊苣株高50-60 cm,留茬5 cm,产量达到50 700 kg/hm^2;高丹草留茬20 cm,产量达到96 300kg/hm^2。     

不同刈割时期和留茬高度对羊草割草地产量和品质的影响

适宜的留茬高度对羊草割草地的牧草生长与牧草营养品质有着重要的作用。本文以呼伦贝尔羊草割草地为研究对象,研究不同的刈割时期与刈割留茬高度对牧草的经济产量及养分含量的影响。本实验设置3个刈割时间,4个留茬高度,每个处理3次重复,采用随机区组实验设计。结果表明,8月15日刈割的植物群落地上生物量显著高于其他刈割时期;随着刈割时间的推迟,群落ADF和NDF含量先上升后下降。留茬高度2 cm的群落地上生物量显著高于留茬高度8 cm(P<0.05),但与留茬高度5 cm间无显著差异(P>0.05),不同刈割时期不同留茬高度下群落粗蛋白(CP)含量无显著差异(P>0.05)。呼伦贝尔羊草割草地适宜刈割时间为8月15日左右,以留茬高度2-5 cm为宜。     

Effects of selective defoliation and height of defoliation on tiller dynamics and herbage yield of Themeda triandra in a semi-arid rangeland in Zimbabwe

Understanding the impacts of intensity and selection of grazing on the performance of grasses is important in the management of grazing areas. It is especially important in semi-arid environments where, apart from moisture, the levels at which grasses are utilized has a major influence on their persistence in the environment. The effects of selective defoliation and height of defoliation (5 cm and 10 cm stubble heights) on the performance of the grass, Themeda triandra, were investigated in a field experiment for two growing seasons. Performance was measured as tiller production, rate of production, tiller mortality, herbage yield and quality. Tiller production was greater (81 tillers per plant) under non-selective defoliation than under selective defoliation (49 tillers per plant) in the 1995/1996 season. Tiller mortality was higher (66.45%) under heavy selective defoliation than under nonselective defoliation (21.98%). Herbage yield, apart from the control treatment, was high (13.6 g per plant) under light non-selective defoliation. Heavy selective defoliation reduced the nutrient levels (e.g. levels of soluble carbohydrates under heavy selective defoliation were 6 g kg^?1 glucose compared to 20 g kg^?1 glucose under light non-selective defoliation).     

Livestock Forage Conditioning Among Six Northern Great Basin Grasses

Studies of Anderson and Scherzinger's forage conditioning hypothesis have generated varied results. Our objectives were: 1) to evaluate late summer/early fall forage quality of crested wheatgrass (Agropyron desertorum [Fisch. ex Link] J. A. Schultes), bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Löve), Idaho fescue (Festuca idahoensis Elmer), bottlebrush squirreltail (Elymus elymoides [Raf.] Swezey), Thurber's needlegrass (Achnatherum thurberianum [Piper] Barkworth), and basin wildrye (Leymus cinereus [Scribn. & Merr.] A. Löve) from ungrazed paddocks and paddocks grazed at vegetative, boot, and anthesis; and 2) test hypotheses that postgrazing regrowth yields were correlated with soil moisture content when grazing occurred. Crop–year precipitation for 1997 and 1998 was 134% and 205% of average. Crude protein (CP) and in vitro dry matter digestibility (IVDMD) of ungrazed grasses displayed expected declines in quality. Among ungrazed grasses, late summer/early fall CP was 5.7% in 1997 and 3.6% in 1998; IVDMD was 47% and 41%, respectively. Late summer/early fall forage quality was elevated by vegetative, boot stage, or anthesis grazing. The phenologically youngest regrowth always ranked highest in CP and IVDMD. Among grasses, respective 1997 CP and IVDMD means were 9.0% and 55% for regrowth following anthesis grazing. No regrowth followed anthesis grazing in 1998, but CP and IVDMD means from boot stage treatments were 5.5% and 47%, respectively. With CP measures, a species by treatment interaction occurred in 1997, but species reacted similarly in 1998. Vegetative, boot stage, and anthesis grazing in 1997 caused respective late summer/early fall standing crop reductions of 34%, 42%, and 58%; and 34%, 54%, and 100% reductions in 1998. Forage conditioning responses were lower for bluebunch wheatgrass and crested wheatgrass than other grasses. Soil moisture content was a poor predictor of regrowth yields. Managed cattle grazing can successfully enhance late season forage quality.     

细胞分裂素调控的不同茬高黑麦草再生机制

通过检测植物生长激素研究不同茬高多花黑麦草(Lolium multiflorum)叶再生的机制。试验结果表明,遮光条件下,高茬和高茬断根的再生叶片生物量均显著高于低茬的再生叶片生物量(P＜0.05)。光照试验结果表明,低茬在单次去叶时、高茬在多次去叶时均易提高总生物量系数(总生物量系数表示单位茬质量所支撑的再生过程中总有机物质量的变化,是用来评价非贮存有机物质参与叶片的再生能力,是不同茬高黑麦草再生能力强弱的指标)。通过检测光照条件下的叶片和伤流液中玉米素+玉米素核苷(Z+ZR)及脱落酸(ABA)含量,发现叶Z+ZR含量与总生物量系数相关密切,且叶Z+ZR含量受根系的直接调控。基于根系对叶片的影响,根系诱导的叶片细胞分裂素是影响不同茬高黑麦草再生的关键因素。     

A Nondestructive Method to Estimate Standing Crop of Purple Threeawn and Blue Grama

We used multiple regression analysis to develop models to predict standing crop of purple threeawn (Aristida purpurea Nutt.) and blue grama (Bouteloua gracilis [H.B.K.] Griffiths) nondestructively. Data were collected for 3 yr on the Texas Tech University Native Rangeland, Lubbock, TX, USA. Independent variables included plant length and area measurements (basal area and cross-sectional area at a 7.5-cm plant height and at 50% of total plant height). One hundred randomly selected plants of each species were measured in June 2008; 50 plants of each species were measured in June 2009 and 2010. Coefficients of determination exceeded 0.91 for both species in all 3 yr of measurement. For both species and years, cross-sectional area at 7.5 cm was the most important single predictor variable. For each species, models differed among years. Our regression models were successful at predicting mid- to late-season standing crop of purple threeawn and blue grama grass and provide an effective method for nondestructive monitoring of these species. This approach should be applicable to similar morphotypes of these species.     

Influence of Plant Functional Group Removal on Inorganic Soil Nitrogen Concentrations in Native Grasslands

High plant functional group diversity has been hypothesized to reduce resource concentrations based on the assumption that species from one functional group acquire resources similarly to one another, while species from other functional groups acquire resources dissimilarly. To determine if functional groups use soil nutrients different from one another, we investigated the impact of removing individual functional groups on soil inorganic nitrogen (NO-3 and NH+4) concentrations in the Idaho fescue (Festuca idahoensis Elmer)/bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Löve) habitat type in Montana. Treatments were imposed by removing 1) all plant species (total plant removal), 2) shallow-rooted (< 15 cm) forbs, 3) deep-rooted (> 15 cm) forbs, 4) all forbs (total forb removal), 5) grasses, and 6) spikemoss, compared to intact control plots. Inorganic nitrogen was measured at 2 soil depths (0–15 cm and 16–40 cm) in the spring, summer, and fall 1 year after treatment imposition. The removal of individual functional groups generally increased soil NO-3 and NH+4 concentrations. Total plant removal increased NO-3 concentrations more than removing individual functional groups. Grass removal generally increased soil NO-3 concentrations in the 0–15-cm depth more than other functional groups removal. Whether the grass or total forb removal treatment resulted in greater soil NH+4 concentrations in the 0–15-cm depth depended on season. These results suggest that functional groups vary in their soil nutrient acquisition patterns and that increased functional diversity decreases soil nutrient concentrations. Therefore, maintaining or improving functional diversity may be a method to more fully utilize soil nutrients because functional groups can differ in their spatial and temporal acquisition of resources.     

Defoliation Effects on Herbage Production and Root Growth of Wet Meadow Forage Species

Root growth is important to the competitive ability of plants, and understanding how herbage defoliation affects root growth has implications for development of management strategies. Objectives were to determine the effects of defoliation intensity and frequency on root characteristics and herbage production of slender wheatgrass (Elymus trachycaulus [Link.] Shinners), Nebraska sedge (Carex nebrascensis C. Dewey), and “Steadfast” birdsfoot trefoil (Lotus corniculatus L.). Plants of each species were transplanted into containers that had been placed in the ground at wet meadow field sites the prior year. There were eight replications of a control and five defoliation treatments, which were combinations of different frequencies (two or five times) and intensities (light or heavy) and haying. Treatments were applied for a single growing season, and aboveground biomass was collected. Containers were extracted in October, and plant crowns, rhizomes, and roots were separated from the soil. Defoliation treatment did not affect total root weight, length, and surface area of Nebraska sedge or birdsfoot trefoil (P>0.10). Slender wheatgrass total root weight was less when defoliated five times (4.46 g·container^?1) than when defoliated twice (6.62 g·container^?1) during the growing season. More frequent defoliation of slender wheatgrass also reduced length (20%) and surface area (21%) compared to less frequent defoliation. However, defoliation frequency did not affect aboveground biomass. Defoliation intensity did not affect aboveground production or root characteristics of the three species. Abundant soil moisture in meadows likely buffers negative effects of defoliation. For all species, two defoliation events (e.g., haying followed by grazing) does not appear to negatively affect root growth and herbage production.     

Root Responses to Short-Lived Pulses of Soil Nutrients and Shoot Defoliation in Seedlings of Three Rangeland Grasses

Root proliferation is important in determining root foraging capability of rangeland grasses to unpredictable soil-nutrient pulses. However, root proliferation responses are often confounded by the inherent relative growth rate (RGR) of the particular species being compared. Additionally, inherent biomass allocation to roots (R:S ratio) can be associated with root RGR, hence likely influencing root foraging responses. The influence of relative growth rate and biomass allocation patterns on the speed and efficiency of root foraging responses at the critical seeding stage was examined in two important perennial rangeland grasses that occur widely in the Great Basin Region of the United States (Whitmar bluebunch wheatgrass [Pseudoroegneria spicata {Pursh} Löve] and Hycrest crested wheatgrass [Agropyron desertorum {Fisch. ex Link} Schult. × A. cristatum L. Gaert.]) as well as in the widespread exotic invasive annual grass, cheatgrass (Bromus tectorum L.). Greenhouse-grown seedlings were exposed to four nutrient regimes: uniform–low, uniform–high, soil-nutrient pulse, soil-nutrient depletion, and to either no clipping or clipping (80% removal of standing shoot biomass). Hycrest was the only species that exhibited root proliferation responses to the short-lived nutrient pulse, and this response occurred through root elongation rather than initiation of lateral root branches. Overall, defoliation inhibited proliferation-based root responses to a larger extent than topological-based root responses. Defoliated plants of Hycrest interrupted root development (topological index did not change) following shoot defoliation compared to undefoliated plants. In contrast, root topological developmental patterns were the same for defoliated and undefoliated plants of Whitmar, whereas cheatgrass exhibited an intermediate response between Whitmar and Hycrest. Our results suggest that inherent biomass allocation to roots contributes to enhanced capabilities of proliferation-based root responses.     

Seedling Defoliation and Drought Stress: Variation in Intensity and Frequency Affect Performance and Survival

Our ability to restore rangelands is limited, and it is unknown if seedling herbivory on its own, or in interaction with other stressors, is a major contributor to restoration failure. To address this, we conducted two experiments: a No Defoliation (ND) experiment (n = 48), in which seedlings from three perennial grasses (crested wheatgrass [Agropyron cristatum {(L.} Gaertn.], bluebunch wheatgrass [Psuedoroegnaria spicata {Pursh} Á. Love], Sandberg bluegrass [Poa secunda J Presl]) were subjected to wet and dry water regimes for 4 mo, and a concurrent Defoliation (D) experiment (n = 95), in which seedlings were factorially assigned to two defoliation treatments—frequency (LOW, HIGH) and intensity (30% vegetation removal, 70% vegetation removal). Indicators of seedling performance were aboveground and belowground biomass (AGB and BGB), root:shoot ratio, tillering, and mortality. The effect size statistic, Hedge’s g, allowed for comparisons between performance measures. Water stress induced reductions in most performance measures: BGB (g = ND: –1.3; D: –1.6), root:shoot ratio (g = ND: n.s.; D: –0.2), and tillering (g = ND: –1.7; D: –1.2), though not significantly for all species. For ABG, water stress interacted with defoliation, reducing performance less at an intensity of 70% (g = –2.0) as opposed to 30% (g = –3.0), but not always significantly in the former. Water stress also caused less reduction in AGB when no defoliation occurred (ND: –0.8; g = D: –2.5). Intensity and frequency of defoliation interacted; seedlings were generally resistant to reductions in performance except at high frequency, 70% defoliation. Agropyron cristatum and P. spicata displayed similar sensitivity to treatments, mostly in terms of changes in AGB and BGB, while P. secunda also experienced increased mortality and reduced tillering. If these differences in sensitivity result in differential survival, herbivory could impact species postrestoration population demographics.