Physiological and ecological constraints play key roles in the evolution of plant growth patterns, especially in relation to defenses against herbivores. Phenotypic and life history theories are unified within the growth-differentiation balance (GDB) framework, forming an integrated system of theories explaining and predicting patterns of plant defense and competitive interactions in ecological and evolutionary time. Plant activity at the cellular level can be classified as growth (cell division and enlargement) of differentiation (chemical and morphological changes leading to cell maturation and specialization). The GDB hypothesis of plant defense is premised upon a physiological trade-off between growth and differentiation processes. The trade-off between growth and defense exists because secondary metabolism and structural reinforcement are physiologically constrained in dividing and enlarging cells, and because they divert resources from the production of new leaf area. Hence the dilemma of plants: Th...

https://www.jstor.org/stable/pdfplus/2830650.pdf

The dilemma of plants: To grow or defend.

Storage is a characteristic feature of most plants, particularly perennials, and the subject has been thoroughly reviewed according to its chemistry and physiology (9, 40, 59, 80, 133, 139). However, in ecology much of the information on storage is based on observation rather than experimentation, and experiments often fail to confirm common perceptions of the nature and dynamics of stored reserves. For example, clipping studies show that not all carbohydrates are available to the plant, even though they are considered to be stored reserves. In this review we suggest criteria for defining storage in ecological and eonomic contexts in order to examine the costs and benefits of storage. We then evaluate the evidence for, and ecological importance of, different types of storage. We discuss storage in relation to vegetative growth

/pdf/the-ecology-and-economics-of-storage-in-plants-2grclrzphb.pdf

The Ecology and Economics of Storage in Plants

Several hypotheses, mainly Optimal Defense (OD), Carbon:Nutrient Balance (CNB), Growth Rate (GR), and Growth‐Differentiation Balance (GDB), have individually served as frameworks for investigating the patterns of plant defense against herbivores, in particular the pattern of constitutive defense. The predictions and tests of these hypotheses have been problematic for a variety of reasons and have led to considerable confusion about the state of the “theory of plant defense.” The primary contribution of the OD hypothesis is that it has served as the main framework for investigation of genotypic expression of plant defense, with the emphasis on allocation cost of defense. The primary contribution of the CNB hypothesis is that it has served as the main framework for investigation of how resources affect phenotypic expression of plant defense, often with studies concerned about allocation cost of defense. The primary contribution of the GR hypothesis is that it explains how intrinsic growth rate of p...

/pdf/out-of-the-quagmire-of-plant-defense-hypotheses-2w74zmn5qu.pdf

Out Of The Quagmire Of Plant Defense Hypotheses

Characteristics of higher plant terpenoids that result in mediation of numerous kinds of ecological interactions are discussed as a framework for this Symposium on Chemical Ecology of Terpenoids. However, the role of terpenoid mixtures, either constitutive or induced, their intraspecific qualitative and quantitative compositional variation, and their dosage-dependent effects are emphasized in subsequent discussions. It is suggested that little previous attention to these characteristics may have contributed to terpenoids having been misrepresented in some chemical defense theories. Selected phytocentric examples of terpenoid interactions are presented: (1) defense against generalist and specialist insect and mammalian herbivores, (2) defense against insect-vectored fungi and potentially pathogenic endophytic fungi, (3) attraction of entomophages and pollinators, (4) allelopathic effects that inhibit seed germination and soil bacteria, and (5) interaction with reactive troposphere gases. The results are integrated by discussing how these terpenoids may be contributing factors in determining some properties of terrestrial plant communities and ecosystems. A terrestrial phytocentric approach is necessitated due to the magnitude and scope of terpenoid interactions. This presentation has a more broadly based ecological perspective than the several excellent recent reviews of the ecological chemistry of terpenoids.

Higher plant terpenoids:a phytocentric overview of their ecological roles

Phenolics have been considered classic defence compounds for protecting plants from herbivores, ever since plant secondary metabolites were suggested to have evolved for that reason. The resource availability and carbon-nutrient balance hypotheses proposed that variation in phenolic levels between and within plant species reflects environmental availability of nutrients and light, and represents a trade-off in allocation by plants to growth and defence against herbivores. In contrast to these concepts, we suggest that (1) the main role of many plant phenolics may be to protect leaves from photodamage, not herbivores; (2) they can achieve this by acting as antioxidants; and (3) their levels may vary with environmental conditions in order to counteract this potential photodamage. We therefore suggest that patterns in phenolic levels, often used to support the concept of trade-off between growth and herbivore defence in relation to resource availability, may actually reflect different risks of photodamage. We suggest that the level of many phenolics is low under some environmental conditions, not because resources to produce them are limited, but simply because the risk of photodamage is low and they are not required. If our photodamage hypothesis is correct, a reassessment of the ecological and evolutionary role of many phenolics in plant defence theory is required.

Rethinking the role of many plant phenolics - protection from photodamage not herbivores?

Plant carbon/nutrient balance has been implicated as an important factor in plant defensive chemistry and palatability to herbivores. We tested this hypothesis by fertilizing juvenile growth form Alaska paper birch and green alder with N, P and N-plus-P in a balanced 2x2 factorial experiment. Additionally, we shaded unfertilized plants of both species. Fertilization with N and N-plus-P increased growth of Alaska paper birch, reduced the concentration of papyriferic acid in internodes and increased the palatability of birch twigs to snowshoe hares. Shading decreased birch growth, decreased the concentration of papyriferic acid in internodes and increased twig palatability. These results indicate that the defensive chemistry and palatability of winter-dormant juvenile Alaska paper birch are sensitive to soil fertility and shade. Conversely the defensive chemistry and palatability of green alder twigs to snowshoe hares were not significantly affected by soil fertility or shade. The greater sensitivity of Alaska paper birch defensive chemistry and palatability to snowshoe hares in comparison to green alder is in agreement with the hypothesis that early successional woody plants that are adapted to high resource availability are more plastic in their chemical responses to the physical environment than are species from less favorable environments.

Response of winter chemical defense in Alaska paper birch and green alder to manipulation of plant carbon/nutrient balance

Palatabilities of parts and growth stages of balsam poplar (Populus balsamifera) to snowshoe hares (Lepus americanus) are related to concentrations of specific plant metabolites that act as antifeedants. Buds are defended from hares by cineol, benzyl alcohol, and (+)-α-bisabolol. Internodes are defended by 6-hydroxycylohexenone (6-HCH) and salicaldehyde. Although defense of interaodes depends upon both compounds, the defense of juvenile internodes is principally related to salicaldehyde concentration; the defense of internode current annual growth is principally related to 6-HCH concentration. The concentration of 6-HCH can be supplemented by the hydrolysis of phenol glycosides when plant tissue is disrupted, raising the possibility of a dynamic element of the chemical defense of poplar.

Winter chemical defense of Alaskan balsam poplar against snowshoe hares.

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Response of winter chemical defense in Alaska paper birch and green alder to manipulation of plant carbon/nutrient balance

Winter chemical defense of Alaskan balsam poplar against snowshoe hares.