The Trials and Tribulations of the Plant Male Gametophyte — Understanding Reproductive Stage Stress Tolerance

Yield and productivity of many crop species depend on successful reproductive development to produce seeds or fruits for human nutrition. Plants determine the right time to flower based on environmental cues (day length, temperature) and angiosperms have evolved a plethora of mechanisms to adapt flowering to specific environmental conditions. Despite these adaptation mechanisms, fertilisation and seed production remain subject to the reigning weather conditions before and during flowering. To fertilise the immobile female gametes inside the ovule, the male gametophytes need to be dispersed in a hostile environment. In crop plants, unexpected inclement weather conditions during male gametophyte development and pollen dispersal are often associated with dramatic yield losses. Molecular and physiological studies are gradually making progress in identifying genes and processes that control various aspects of pollen development, but the many intricacies involved in environmental control of pollen development and – in particular – regulation of male fertility remain poorly understood. The aim of this paper is to draw attention to the enormous amount of complexity and biodiversity that exist in angiosperm male gametophyte development. A better understanding of the strategies that exist in adapting pollen production and fertility to environmental challenges may ultimately benefit improvement of abiotic stress tolerance in major food crops

Inter-Conversion of carbohydrate reserves from pollen maturation to rehydration in a chili pepper

Carbohydrate metabolism is critical for male fertility. Carbohydrate reserves (soluble sugars and starch) were quantified in three advanced stages of pollen development, from the final maturation to rehydration, in a chili pepper cultivar (Capsicum annuum L. cv. Calypso Red) with partially dehydrated pollen, to define possible carbohydrate inter-conversions. At the same time, the activity of soluble enzymes involved in sucrolysis and amylolysis were quantified to reveal potential metabolic pathways. The carbohydrates found (sucrose, glucose, fructose, and starch) apparently inter-convert. There would be a close relation between enzymatic activities and substrates, and the carbohydrate reserves could be key factors on the regulation of enzymatic activities. All the enzymes tested were active, although the relevance of each one changed along pollen maturation and rehydration, defining different profiles of enzymatic activities for each stage. Some usually neglected enzymes (e.g. soluble neutral invertase) have shown an important role in the stages analyzed, suggesting alternative processes to evaluate in the studies of male fertility control.Fil: Carrizo Garcia, Carolina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Córdoba. Instituto Multidisciplinario de Biología Vegetal (p); ArgentinaFil: Guarnieri, Massimo. Universita Degli Studi Di Siena; ItaliaFil: Pacini, Ettore. Universita Degli Studi Di Siena; Itali

Pollen Developmental Arrest: Maintaining Pollen Fertility in a World With a Changing Climate

During evolution of land plants, the haploid gametophytic stage has been strongly reduced in size and the diploid sporophytic phase has become the dominant growth form. Both male and female gametophytes are parasitic to the sporophyte and reside in separate parts of the flower located either on the same plant or on different plants. For fertilization to occur, bi-cellular or tri-cellular male gametophytes (pollen grains) have to travel to the immobile female gametophyte in the ovary. To survive exposure to a hostile atmosphere, pollen grains are thought to enter a state of complete or partial developmental arrest (DA). DA in pollen is strongly associated with acquisition of desiccation tolerance (DT) to extend pollen viability during air travel, but occurrence of DA in pollen is both species-dependent and at the same time strongly dependent on the reigning environmental conditions at the time of dispersal. Several environmental stresses (heat, drought, cold, humidity) are known to affect pollen production and viability. Climate change is also posing a serious threat to plant reproductive behavior and crop productivity. It is therefore timely to gain a better understanding of how DA and pollen viability are controlled in plants and how pollen viability can be protected to secure crop yields in a changing environment. Here, we provide an overview of how DA and pollen viability are controlled and how the environment affects them. We make emphasis on what is known and areas where a deeper understanding is needed

Fruit and seed structural characteristics and seed dispersal in Mercurialis annua L. (Euphorbiaceae)

The fruit of Mercurialis annua L. is a two-seeded capsule with a caruncle, a small appendage which arises from the outer integument in the micropylar area. The inner integument has a thick layer of Malpighian-like cells, interrupted at the micropyle by a remnant of nucellus, the walls of which contain suberin. There is no cuticle covering the caruncle and its cells have lipids and protein bodies as reserves. Seed dispersal occurs by a combination of autochory followed by myrmecochory. The explosive ejection of seeds is caused by dehydration-induced torsion of the mechanical layer of the fruit walls. This phenomenon is due to the different orientation of the cells and a folding over of the mechanical layer in the chalazal area. The seeds were found to be thrown distances from 1 to 130 cm (mean 41.1 cm; modal peak 10-20 cm). Ballistic dispersal is influenced by the weight of the seed. The seeds were collected by ants in a mean time of 24.4 minutes the maximum and mean distances of dispersal being 14 and 3.4 m, respectively. Of the seeds removed, 95.2 % were recovered on excavation of the ant nest. The seeds in the nest were intact but without the caruncles. The mean distance of dispersal by diplochory was < 5 m. These features are discussed in relation to possible benefits to the plant species in terms of adaptive advantages of seeds

Some cytological, ecological and evolutionary aspects of pollination

Different types of pollen dispersing units are described. The pollen of angiosperms may be dispersed in monads, tetrads, polyads, massulae or compact pollinia. The monads and tetrads may form larger clumps of pollen with the aid of the following substances: a) pollenkitt, a product of in situ degeneration of the tapetum; b) tryphine, a product of tapetal extra situm degeneration, found only in Cruciferae; c) elastoviscin, a highly viscous product of the degeneration of a limited number of tapetal cells, found only in certain Orchidaceae; d) viscin threads or sporopollenin filaments arising from the pollen exine. The number of pollen grains composing the clumps depends on pollen diameter and the cohesive forces holding the pollen grains together. The various types of pollen dispersing unit are discussed in relation to the type of pollination and number of ovules per ovary

Pollen grain sporoderm and types of dispersal units

The pollen of gymnosperms and angiosperms may be dispersed in monads, tetrads, polyads, massulae or compact pollinia. The monads and tetrads may form larger clumps of pollen because filiform pollen is tangled while other kinds of pollen can be glued by means of different devices. Exine and intine modify their structure to adapt to pollen dispersing units, exine in some cases can be absent. An additional layer, a thin callosic wall, can be present in some species beneath the intine; this occurs when pollen grains are slightly dehydrated before dispersal

Ultrastructural Aspects of Unilateral Interspecific Incompatibility between Lycopersicum Peruvianum and L. Esculentum

SUMMARYObservations have been made, at the electron microscope, of the pollen tubes present in the styles of Lycopersicum esculentum and L. peruvianum after reciprocal crosses between the two species.The unilateral incompatibility barrier which isolates the two species when L. peruvianum is used as pistillate parent was then compared to the processes of pollen tube rejection which have been recently analysed (J. Cell Sci., 1972) after self-pollination in this self-incompatible species. Such a comparison, which was also carried out by means of fluorescence techniques, has permitted to find out that for both types of incompatibility the rejection process was characterised by a progressive disappearance of the callose-rich inner wall of the pollen tube and by an accumulation of bi-partite particles in the tube cytoplasm.In the case of unilateral incompatibility, however, the tube outer wall is gradually disaggregated while the callosic inner wall remains quite thick at the tube apex, becoming thinner and fin..

An unexplored side of regeneration niche: seed quantity and quality are determined by the effect of temperature on pollen performance

In 1977, Peter Grubb introduced the regeneration niche concept, which assumes that a plant species cannot persist if the environmental conditions are only suitable for adult plant growth and survival, but not for seed production, dispersal, germination and seedling establishment. During the last decade, this concept has received considerable research attention as it helps to better understand community assembly, population dynamics, and plant responses to environmental changes. Yet, in its present form, it focuses too much on the post-fertilization stages of plant sexual reproduction, neglecting the fact that the environment can operate as a constraint at many points in the chain of processes necessary for successful regeneration. In this review, we draw the attention of the plant ecology research community to the pre-fertilization stages of plant sexual reproduction, an almost ignored but important aspect of the regeneration niche, and their potential consequences for successful seed production. Particularly, we focus on how temperature effects pollen performance and determines plant reproduction success by playing an important role in the temporal and spatial variations in seed quality and quantity. We also review the pollen adaptations to temperature stresses at different levels of plant organization and discuss the plasticity of the performance of pollen under changing temperature conditions. The reviewed literature demonstrates that pre-fertilization stages of seed production, particularly the extreme sensitivity of male gametophyte performance to temperature, are the key determinants of a species’ regeneration niche. Thus, we suggest that previous views stating that the regeneration niche begins with the production of seeds should be modified to include the preceding stages. Lastly, we identify several gaps in pollen-related studies revealing a framework of opportunities for future research, particularly how these findings could be used in the field of plant biology and ecology

Polyphenol-rich diets improve glucose metabolism in people at high cardiometabolic risk: a controlled randomised intervention trial

Dietary polyphenols and long chain n-3 polyunsaturated fatty acids (LCn3) are associated with lower cardiovascular risk. This may relate to their influence on glucose metabolism and diabetes risk. We evaluated the effects of diets naturally rich in polyphenols and/or LCn3 of marine origin on glucose metabolism in people at high cardiometabolic risk

In search of traces of the mandrake myth: the historical, and ethnobotanical roots of its vernacular names

Mandrake (Mandragora spp.) is one of the most famous medicinal plant in western cultures since Biblical times and throughout written history. In many cultures, mandrake is related to magic and witchcraft, which is said to have a psychosomatic effect (especially when mandrake contains narcotic compounds) in addition to the pharmacological influence, as occurs with other narcotic magical plants. Due to its unique properties and related myths, it is not surprising that this plant has many names in many languages. Methods: This paper presents an attempt to reconstruct the historical, ethnobotanical, and folkloristic roots of 292 vernacular names of Mandragora spp. in forty-one languages. We used the plant’s morphological data, philology, myths and legends, medicinal properties and uses, as well as historical evidence and folkloric data, to explain meaning, origin, migration, and history of the plant’s names. Results: The names were classified into the following main categories: Derivatives of mandragora (19 languages), alraun (7) and of yabroukh (5). The salient groups of the plant’s vernacular names are related to: Anthropomorphism (33 names in 13 languages); Similarity to other plants (28/9); Supernatural agents (28/9); Narcotic effects (21/8); Leaves, fruits, and seeds (21/8); Aphrodisiac properties (17/10); Use of a dog (15/9); Gallows (14/5); Black magic, sorcery, witchcraft (13/8), and Medicinal use (11/7). Conclusions: This frequency distribution of the mandrake’s vernacular names reflects its widespread reputation as related to the doctrine of signatures, beliefs in its supernatural, natural, and mythic powers, and to a lesser extent, its uses in magic and medicine. A spatiotemporal analysis of the mandrake’s names supports the old idea that the pulling ceremonies for this plant originated in the Near East and that various other myths related to this plant may have originated in different places and periods