Leaf elongation rate (LER) in grasses is dependent on epidermal cell supply (number) and on rate and duration of epidermal cell elongation. Nitrogen (N) fertilization increases LER. Longitudinal sections from two genotypes of tall fescue (Festuca arundinacea Schreb.), which differ by 50% in LER, were used to quantify the effects of N on the components of epidermal cell elongation and on mesophyll cell division. Rate and duration of epidermal cell elongation were determined by using a relationship between cell length and displacement velocity derived from the continuity equation. Rate of epidermal cell elongation was exponential. Relative rates of epidermal cell elongation increased by 9% with high N, even though high N increased LER by 89%. Duration of cell elongation was approximately 20 h longer in the high-than in the low-LER genotype regardless of N treatment. The percentage of mesophyll cells in division was greater in the high-than in the low-LER genotype. This increased with high N in both genotypes, indicating that LER increased with cell supply. Division of mesophyll cells adjacent to abaxial epidermal cells continued after epidermal cell division stopped, until epidermal cells had elongated to a mean length of 40 micrometers in the high-LER and a mean length of 50 micrometers in the low-LER genotype. The cell cycle length for mesophyll cells was calculated to be 12 to 13 hours. Nitrogen increased mesophyll cell number more than epidermal cell number: in both genotypes, the final number of mesophyll cells adjacent to each abaxial epidermal cell was 10 with low N and 14 with high N. A spatial model is used to describe three cell development processes relevant to leaf growth. It illustrates the overlap of mesophyll cell division and epidermal cell elongation, and the transition from epidermal cell elongation to secondary cell wall deposition.Leaf growth in grasses is predominantly unidirectional, parallel with the longitudinal axis of the leaf. A 3 Abbreviations: SLW, specific leaf weight; LER, leaf elongation rate; PPFD, photosynthetic photon flux density. 549Volenec and Nelson (29) used high and low rates of N fertilization to further alter LER of two genotypes of tall fescue selected for contrasting leaf growth rates. Mean LER of these genotypes was 89% higher with high N, but length of fully elongated epidermal cells was unaffected by N treatment. The number of epidermal cells produced per file per day increased 90% with high N, suggesting that much of the increase in LER following N fertilization was due to increased cell division.Our long-term goal is to understand carbohydrate metabolism during the growth of grass leaves. Our objectives in the present study were to evaluate the effect of N on mesophyll cell division and rate and duration of epidermal cell elongation in leaf blades of two genotypes of tall fescue. Data for epidermal cell lengths utilized here have been reported previously (28, 29).Mesophyll cell division data reported here, and data for leaf elongation rate and structural ...

Whether foraging on pastures or rangelands, herbivores encounter plant species that differ in their concentrations of nutrients. They also all contain various secondary compounds that at too high doses can be toxic, but at the appropriate dose many of these toxins may have medicinal benefits. The quantity of forage an animal consumes depends on the other forages it selects because nutrients and toxins interact. Food intake also depends on an individual's morphology and physiology, and marked variation is common, even among closely related animals, in needs for nutrients and abilities to cope with toxins. Thus, individuals can better meet their needs when offered a variety of foods that differ in nutrients and toxins than when constrained to a single food. Nonetheless, we have focused on a few species, often grown in monoculture, and we have reduced concentrations of secondary compounds with little appreciation for their roles in protecting plants against herbivores, pathogens, and competitors. In nature, where diversity of plants is the rule and not the exception, eating a variety of foods is how animals cope with, and may benefit from, secondary compounds. The potential benefits of creating mixtures of plant species whose nutrient and secondary compound profiles complement one another are obvious, though much remains to be learned about how to reconstruct agro‐ecosystems with plants that complement and enhance one another structurally, functionally, and biochemically.

Cessation of cell expansion has been associated with cell wall cross-linking reactions catalyzed by peroxidase. This study utilized two genotypes of tall fescue (Festuca arundinacea Schreb.) that differ in length of the leaf elongation zone to investigate the relationship between ionically bound peroxidase activity and the spatial distribution of leaf elongation. Peroxidase activity was also localized histochemically in transverse sections of the leaf blade using 3,3'-diaminobenzidine. Soluble or soluble plus ionically bound peroxidase activities were extracted from homogenized segments of the elongating leaf blade and assayed spectrophotometrically. Activity of the ionically bound fraction, expressed per milligram fresh weight or per microgram protein, increased as cells were displaced through the distal half of the elongation zone, corresponding to the region in which the elongation rate declined. In both genotypes, the initial increase in activity preceded the onset of growth deceleration by about 10 hours. In the basal region where elongation began, histochemical localization showed that peroxidase activity was found only in vascular tissues. As cells were displaced farther through the elongation zone, peroxidase activity appeared in walls of other longitudinally continuous tissues such as the epidermis and bundle sheaths. Increase in ionically bound peroxidase activity and changes in localization of peroxidase activity occurred at comparable developmental stages in the two genotypes. The results indicate that cessation of elongation followed an increase in cell wall peroxidase activity.

We examined relationships among cell wall feruloylation, diferulate cross-linking, p-coumarate deposition, and apoplastic peroxidase (EC 1.11.1.7) activity with changes in the elongation rate of leaf blades of slow and rapid elongating genotypes of tall fescue ( Festuca arundinacea Schreb.). Growth was not directly influenced by ferulic acid deposition but leaf elongation decelerated as 8-5-, 8- O-4-, 8-8-, and 5-5-coupled diferulic acids accumulated in cell walls. Growth rapidly slowed and stopped with the deposition of p-coumarate, which is primarily associated with lignification in grass cell walls. Accretion of ferulate, diferulates and p-coumarate continued after growth ended, into the later stages of secondary wall formation. The concentration of 8-coupled diferulates dwarfed that of the more commonly measured 5-5-coupled isomer, suggesting that the latter dimer is a poor indicator of diferulate cross-linking in cell walls. Further work is required to clearly demonstrate the role of diferulate cross-linking and p-coumaroylated lignins in the cessation of leaf growth in grasses.

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The objective of this study was to determine whether feeding tannin-containing hays to heifers and mature beef cows influences enteric methane (CH4) emissions and nitrogen (N) excretion relative to feeding traditional legume and grass hays. Fifteen mature beef cows (Exp. 1) and 9 yearling heifers (Exp. 2) were each randomly assigned to treatment groups in an incomplete bock design with 2 periods and 6 types of hays with 3 hays fed each period (n = 5 cows and 3 heifers per treatment). Groups were fed tannin-containing [birdsfoot trefoil (BFT), sainfoin (SAN), small burnet (SML)] or non-tannin-containing [alfalfa (ALF), cicer milkvetch (CMV), meadow bromegrass (MB)] hays. Each period consisted of 14 d of adjustment followed by 5 d of sample collection. Nine cows and 9 heifers were selected for the measurement of enteric CH4 emissions (sulfur hexafluoride tracer gas technique), and excretion of feces and urine, while dry matter intake (DMI) was measured for all animals. The concentration of condensed tannins in SAN and BFT was 2.5 ± 0.50% and 0.6 ± 0.09% of dry matter (DM), respectively, while SML contained hydrolyzable tannins (4.5 ± 0.55% of DM). Cows and heifers fed tannin-containing hays excreted less urinary urea N (g/d; P < 0.001) and showed lower concentrations of blood urea N (mg/dL; P < 0.001) than animals fed ALF or CMV, indicating that tannins led to a shift in route of N excretion from urine to feces. Additionally, cows fed either BFT or CMV showed the greatest percentage of retained N (P < 0.001). Enteric CH4 yield (g/kg of DMI) from heifers (P = 0.089) was greatest for MB, while daily CH4 production (g/d) from heifers (P = 0.054) was least for SML. However, digestibility of crude protein was reduced for cows (P < 0.001) and heifers (P < 0.001) consuming SML. The results suggest that tannin-containing hays have the potential to reduce urinary urea N excretion, increase N retention, and reduce enteric CH4 emissions from beef cattle. The non-bloating tannin-free legume CMV may also reduce environmental impacts relative to ALF and MB hays by reducing N excretion in urine and increasing N retention.

In irrigated pastures of the semiarid, high‐elevation western USA, perennial ryegrass (Lolium perenne L.) persistence is poor, and over time white clover (Trifolium repens L.) often dominates mixtures. Irrigation is often not available during autumn, when these perennial plants store carbohydrate reserves for spring regrowth. Our objective was to compare the effect of water stress on growth, carbohydrates, and soil water use of perennial ryegrass, white clover, and tall fescue (Festuca arundinacea Schreb.) in a greenhouse study. These three species were grown separately in a Kidman fine sandy loam, in 15‐cm‐diam, 1‐m‐deep pots and irrigated for 81 d (4 plants/pot). Paired pots were then either irrigated or subjected to water deficit (drought) for 30 d, followed by 10 d of recovery with irrigation. At 10‐d intervals, four paired pots of each species were destructively sampled to determine leaf and storage organ dry matter and carbohydrate and simple sugar concentrations in storage organs. Root length density and soil water content were also sampled at 20‐, 60‐, and 90‐cm soil depths. Leaf dry matter was lower in water‐stressed plants than in irrigated plants by the end of the drought, but did not differ among species. After 10 d of recovery, storage carbohydrate concentration in droughted perennial ryegrass was lower than in white clover, and the ratio of simple sugars (droughted:irrigated) in perennial ryegrass was higher than in white clover. Tall fescue performed similarly to both species. Before the drought, grasses had similar, extensive root systems that withdrew more soil water from the 90‐cm soil depth than did white clover. By the end of the 30‐d drought, white clover had reduced soil water at all depths as much as the grasses. White clover survived drought and conserved carbohydrate reserves after 10 d of recovery better than did perennial ryegrass and similarly to tall fescue.

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