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Woody plants such as trees have a significant economic and climatic influence on global economies and ecologies. This completely revised classic book is an up-to-date synthesis of the intensive research devoted to woody plants published in the second edition, with additional important aspects from the authors' previous book, Growth Control in Woody Plants. Intended primarily as a reference for researchers, the interdisciplinary nature of the book makes it useful to a broad range of scientists and researchers from agroforesters, agronomists, and arborists to plant pathologists and soil scientists. This third edition provides crutial updates to many chapters, including: responses of plants to elevated CO2; the process and regulation of cambial growth; photoinhibition and photoprotection of photosynthesis; nitrogen metabolism and internal recycling, and more. Revised chapters focus on emerging discoveries of the patterns and processes of woody plant physiology. * The only book to provide recommendations for the use of specific management practices and experimental procedures and equipment*Updated coverage of nearly all topics of interest to woody plant physiologists* Extensive revisions of chapters relating to key processes in growth, photosynthesis, and water relations* More than 500 new references * Examples of molecular-level evidence incorporated in discussion of the role of expansion proteins in plant growth; mechanism of ATP production by coupling factor in photosynthesis; the role of cellulose synthase in cell wall construction; structure-function relationships for aquaporin proteins

1. Changing winter climate extremes are expected to result in the poleward migration of mangrove forests at the expense of salt marshes. Although mangroves and marshes are both highly valued ecosystems, the ecological implications of mangrove expansion have not been fully investigated. 2. Here, we examined the effects of mangrove expansion on below-ground properties related to peat development and carbon storage. We investigated plant–soil interactions in marshes and across mangrove forest structural gradients in three locations in the northern Gulf of Mexico (USA). We compared our results to those from terrestrial grasslands where the effects of woody plant encroachment are often influenced by rainfall and plant traits. 3. Abiotic conditions at our study locations differed, particularly in terms of physicochemical properties related to precipitation. Marsh species composition, marsh above-ground biomass, and mangrove forest structural complexity also varied across these locations. Marshes in the driest location (Central Texas) had higher salinities and were dominated by low biomass succulent plants and lower soil carbon pools. Marshes in the wetter, less saline locations (Louisiana and North Florida) contained high biomass grasses and higher soil carbon pools. 4. At all locations, above-ground biomass and above-ground carbon pools were higher in mangroves than marshes; however, below-ground soil carbon pools were only higher in mangroves than marshes in the driest location. In the wetter locations, the linkages between mangrove forest structure and soil properties were minimal or not significant. However, in the driest location, there was a significant increase in soil properties related to peat development and carbon storage with increased mangrove forest structural development. 5. Synthesis-. Our results indicate that the ecological implications of woody plant encroachment in tidal saline wetlands are dependent upon precipitation controls of plant–soil interactions. Although the above-ground effects of mangrove expansion are consistently large, below-ground influences of mangrove expansion appear to be greatest along low-rainfall coasts where salinities are high and marshes being replaced are carbon poor and dominated by succulent plants. Collectively, these findings complement those from terrestrial ecosystems and reinforce the importance of considering rainfall and plant–soil interactions within predictions of the ecological effects of woody plant encroachment.

Aim: Disturbances such as fire operate against a backdrop of constraints imposed by climate and soils to influence grass–woody plant abundance. However, little is known of how these factors interact to determine the upper limits of woody cover and stature in grasslands, in which shrub/tree abundance has been increasing globally. Location: Kansas, Oklahoma, Texas, USA. Time period: 2004–2014. Major taxa studied: Angiosperms and gymnosperms. Methods: Using a database of 1,466 sites and quantile regression to derive precipitation-based upper limits to woody cover and height within grasslands of the central/southern Great Plains, USA, we assessed how soil texture and climate-related fire probabilities [two groups; low fire probability, P(Flow), versus high fire probability, P(Fhi)] might influence realization of the climate potential. Results: Soil texture had no substantive influence on regional-scale woody cover, but taller plants were predicted on sandy soils. Woody plant height potential increased linearly with increasing annual precipitation, becoming asymptotic at c. 800 mm for both the P(Flow) and the P(Fhi) fire groups, after which P(Flow) areas were predicted to support taller plants. Potential woody cover also increased linearly with annual precipitation until c. 800 mm, after which predictions of maximum % cover were similar under both fire groups. Main conclusions: Precipitation was the overriding factor constraining potential woody cover and height, particularly in drier regions, with fire playing a minor role at these regional scales. In contrast to height potential, cover potential remained similar for both P(Flow) and P(Fhi) sites. Dynamic adjustments in woody plant architecture and allocation to foliage and stems, wherein areal cover is maintained when height is suppressed has implications for remote sensing, primary production and biogeochemical processes. Our analyses indicate drier grasslands [< 800 mm mean annual precipitation (MAP)] undergoing woody plant encroachment have the potential to become shrublands (e.g. short woody plants, low cover), whereas wetter areas have the potential to become woodland or forest (taller woody plants, high cover).

The vegetation of semi‐arid and arid landscapes is often comprised of mixtures of herbaceous and woody vegetation. Since the early 1900s, shifts from herbaceous to woody plant dominance, termed woody plant encroachment and widely regarded as a state change, have occurred world‐wide. This shift presents challenges to the conservation of grassland and savanna ecosystems and to animal production in commercial ranching systems and pastoral societies. Dryland management focused on cattle and sheep grazing has historically attempted to reduce the abundance of encroaching woody vegetation (hereafter, ‘brush management’) with the intent of reversing declines in forage production, stream flow or groundwater recharge. Here, we assess the known and potential consequences of brush management actions, both positive and negative, on a broader suite of ecosystem services, the scientific challenges to quantifying these services and the trade‐offs among them. Our synthesis suggests that despite considerable investments accompanying the application of brush management practices, the recovery of key ecosystem services may be short‐lived or absent. However, in the absence of such interventions, those and other ecosystem services may be compromised, and the persistence of grassland and savanna ecosystem types and their endemic plants and animals threatened. Addressing the challenges posed by woody plant encroachment will require integrated management systems using diverse theoretical principles to design the type, timing and spatial arrangement of initial management actions and follow‐up treatments. These management activities will need to balance cultural traditions and preferences, socio‐economic constraints and potentially competing land‐use objectives. Synthesis. Our ability to predict ecosystem responses to management aimed at recovering ecosystem services where grasslands and savannas have been invaded by native or exotic woody plants is limited for many attributes (e.g. primary production, land surface–atmosphere interactions, biodiversity conservation) and inconsistent for others (e.g. forage production, herbaceous diversity, water quality/quantity, soil erosion, carbon sequestration). The ecological community is challenged with generating robust information about the response of ecosystem services and their interactions if we are to position land managers and policymakers to make objective, science‐based decisions regarding the many trade‐offs and competing objectives for the conservation and dynamic management of grasslands and savannas.

As the atmosphere warms, precipitation events become larger, but less frequent. Yet, there is fundamental disagreement about how increased precipitation intensity will affect vegetation. Walter’s two-layer hypothesis and experiments testing it have demonstrated that precipitation intensity can increase woody plant growth. Observational studies have found the opposite pattern. Not only are the patterns contradictory, but inference is largely limited to grasslands and savannas. We tested the effects of increased precipitation intensity in a shrubsteppe ecosystem that receives >30% of its precipitation as snow. We used 11 (8 × 8 m) shelters to collect and redeposit rain and snow as larger, more intense events. Total annual precipitation was the same in all plots, but each plot received different precipitation event sizes ranging from 1 to 18 mm. Over three growing seasons, larger precipitation event sizes increased soil water availability, sagebrush (Artemisia tridentata) stem radius, and canopy greenness, decreased new root growth in shallow soils, and had no effect on herbaceous plant cover. Thus, we found that increased precipitation intensity can increase soil water availability and woody plant growth in a cold semiarid system. Assuming that stem growth is positively correlated with shrub reproduction, establishment and spread, results suggest that increasing precipitation intensity may have contributed to the woody plant encroachment observed around the world in the past 50 yr. Further, continuing increases in precipitation intensity caused by atmospheric warming are likely to continue to contribute to shrub encroachment in the future.

For many woody species, such as Quercus robur, cytokinins in the culture medium are required to maintain in vitro plant material. Among synthetic cytokinins, 6-benzylaminopurine (BAP) and kinetin (KIN) are the most frequently used. In addition to inducing shoots, cytokinins can cause morphophysiological disorders. Therefore, we aimed to investigate the anatomical, biochemical, and physiological alterations and profiles of Q. robur shoots exposed to two cytokinins, applied alone and in combination. Shoots previously established in vitro were transferred to WPM culture media supplemented with BAP at concentrations of 0, 1.25, and 3.50 µM combined with KIN at concentrations of 0, 0.62, and 1.25 µM totaling 9 treatments. Anatomical, physiological, and biochemical analyses were performed after 40 days of culture. BAP induced the formation of new buds with anatomically underdeveloped leaves; induced shoot-tip necrosis, which is considered a response to the inefficient transport of water and nutrients; reduced the thickness of the cell walls of phloem fibers; and decreased the content of phenolic compounds and photosynthetic pigments. These responses were less pronounced with co-exposure to KIN. In contrast, KIN alone stimulated a larger area of secondary xylem and more lignified cell walls. BAP can induce shoots with underdeveloped anatomical and biochemical characteristics. Shoots that grew with KIN alone had stem and leaf anatomical characteristics, indicating greater commitment to cellular differentiation than proliferation. When both cytokinins are combined, KIN can partially mitigate the deleterious effects of BAP on in vitro growth.Key messageBAP generated shoots with underdeveloped anatomical and biochemical characteristics. KIN can partially mitigate the deleterious effects of BAP on in vitro growth.

Woody-plant encroachment is a global phenomenon that has been affecting the southwestern United States since the late 1800s. Drought, overgrazing, herbivory, and competition between grasses and shrub seedlings have been hypothesized as the main drivers of shrub establishment. However, there is limited knowledge about the interactions among these drivers. Using a rainfall manipulation system and various herbivore exclosures, we tested hypotheses about how precipitation (PPT), competition between grasses and shrub seedlings, and predation affect the germination and first-year survival of mesquite (Prosopis glandulosa), a shrub that has encroached in Southern Great Plains and Chihuahuan Desert grasslands. We found that mesquite germination and survival (1) increased with increasing PPT, then saturated at about the mean growing season PPT level, (2) that competition between grasses and shrub seedlings had no effect on either germination or survival, and (3) that herbivory by small mammals decreased seedling establishment and survival, while ant granivory showed no effect. In addition to its direct positive effect on survival, PPT had an indirect negative effect via increasing small mammal activity. Current models predict a decrease in PPT in the southwestern United States with increased frequency of extreme events. The non-linear nature of PPT effects on Mesquite recruitment suggests asymmetric responses, wherein drought has a relatively greater negative effect than the positive effect of wet years. Indirect effects of PPT, through its effects on small mammal abundance, highlight the importance of accounting for interactions between biotic and abiotic drivers of shrub encroachment. This study provides quantitative basis for developing tools that can inform effective shrub management strategies in grasslands and savannas.

Evidence suggests that arbuscular mycorrhizal fungi (AMF) may promote the growth of woody plants. However, the effects of AMF on nitrogen (N) metabolism in plants, especially trees, and its regulatory mechanism are rarely reported. Here, the effects of AMF inoculation on the growth and N nutrition status of Catalpa bungei under different N levels were reported. Three N levels (low, medium, high) and two mycorrhizal inoculation treatments (inoculation with Rhizophagus intraradices or not) were used with factorial design. The results showed that medium N could significantly improve the physiological metabolism and growth of C . bungei seedlings. However, when N was excessive, growth was significantly inhibited whether inoculated AMF or not. Compared with non-inoculated treatments, AMF inoculation could promote the absorption of N and P, improve photosynthesis under low to medium N levels, thus promoting the growth of seedlings. AMF changed the biomass allocation in seedlings by reducing the stem mass ratio and root/shoot ratio, and increasing the leaf mass ratio. At medium N levels, compared with non-inoculated treatment, AMF inoculation could significantly promote root growth by changing root hormone levels and improving root architecture and root activity. Under N addition, AMF inoculation could improve the absorption and assimilation of N by regulating the expression of key enzyme genes of N metabolism and nitrate transporter genes ( NRT2.4 , NRT2.5 , NRT2.7 ) in roots, and enhancing the activities of the key enzyme of N metabolism. This study may provide a reference for the application of AMF in the cultivation and afforestation technology of C. bungei in Northwest China.

As the global population increases and the economy grows rapidly, the demand for livestock products such as meat, egg and milk continue to increase. The shortage of feed in livestock production is a worldwide problem restricting the development of the animal industry. Natural woody plants are widely distributed and have a huge biomass yield. The fresh leaves and branches of some woody plants are rich in nutrients such as proteins, amino acids, vitamins and minerals and can be used to produce storage feed such as silage for livestock. Therefore, the development and utilization of natural woody plants for clean fermented feed is important for the sustainable production of livestock product. This paper presents a comprehensive review of the research progress, current status and development prospects of forageable natural woody plant feed resources. The nutritional composition and uses of natural woody plants, the main factors affecting the fermentation of woody plant silage and the interaction mechanism between microbial co-occurrence network and secondary metabolite are reviewed. Various preparation technologies for clean fermentation of woody plant silage were summarized comprehensively, which provided a sustainable production mode for improving the production efficiency of livestock and producing high-quality livestock product. Therefore, woody plants play an increasingly important role as a potential natural feed resource in alleviating feed shortage and promoting sustainable development of livestock product.

1. Recent global trends of increasing woody plant abundance in grass-dominated ecosystems may substantially enhance soil organic carbon (SOC) storage and could represent a strong carbon (C) sink in the terrestrial environment. However, few studies have quantitatively addressed the influence of spatial heterogeneity of vegetation and soil properties on SOC storage at the landscape scale. In addition, most studies assessing SOC response to woody encroachment consider only surface soils, and have not explicitly assessed the extent to which deeper portions of the soil profile may be sequestering C. 2. We quantified the direction, magnitude and pattern of spatial heterogeneity of SOC in the upper 1-2 m of the profile following woody encroachment via spatially specific intensive soil sampling across a landscape in a subtropical savanna in the Rio Grande Plains, USA, that has undergone woody proliferation during the past century. 3. Increased SOC accumulation following woody encroachment was observed to considerable depth, albeit at reduced magnitudes in deeper portions of the profile. Overall, woody clusters and groves accumulated 12-87 and 18-67 Mg ha⁻¹ more SOC compared to grasslands to a depth of 1·2 m. 4. Woody encroachment significantly altered the pattern of spatial heterogeneity of SOC to a depth of 5 cm, with marginal effect at 5-15 cm, and no significant impact on soils below 15 cm. Fine root density explained greater variability of SOC in the upper 15 cm, while a combination of fine root density and soil clay content accounted for more of the variation in SOC in soils below 15 cm across this landscape. 5. Synthesis. Substantial soil organic carbon sequestration can occur in deeper portions of the soil profile following woody encroachment. Furthermore, vegetation patterns and soil properties influenced the spatial heterogeneity and uncertainty of soil organic carbon in this landscape, highlighting the need for spatially specific sampling that can characterize this variability and enable scaling and modelling. Given the geographic extent of woody encroachment on a global scale, this undocumented deep soil carbon sequestration suggests this vegetation change may play a more significant role in regional and global carbon sequestration than previously thought.