PLANT MORPHOLOGY - T. A. Sautkina - 2012

INTRODUCTION. A BRIEF HISTORY OF THE DEVELOPMENT OF BOTANY

Plant Morphology—the science of structural patterns, processes of morphogenesis, as well as individual and evolutionary Development of Plants—is one of the most fundamental branches of botany. The Development of botany as a science began with humanity's first encounters with higher plants.

Higher plants belong to the kingdom Vegetabilia, or Рlаntае (Plants). Their origin is presumably associated with the transition of unknown ancient Algae onto land. In The process of adapting to terrestrial life, higher plants developed traits that significantly set them apart from their algal ancestors and markedly elevated their level of structural Organization.

Higher plants are characterized by a specialized cellular Structure, the presence of Tissues performing various Functions, and the Differentiation of the body into vegetative and reproductive Organs, which evolved as an adaptation to terrestrial environments. Their reproductive organs are always multicellular, featuring a more advanced sexual process than that of algae; moreover, in highly organized plants—gymnosperms and angiosperms—Fertilization occurs without The Need for an aqueous liquid medium. Finally, higher plants exhibit highly complex life cycles and, most importantly, the vast majority of them are dominated by the sporophyte, which is genetically and functionally better adapted to terrestrial existence than the gametophyte.

The development of higher plants is a complex, multi-stage process that can be conventionally divided into two periods: the first is The formation of the vegetative sphere, and the second is the Formation of the reproductive sphere.

Thanks to vegetative organs, which perform the functions of soil and aerial Nutrition, Transpiration, gas exchange, and the secretion of metabolic products, the individual life of each Organism is sustained. At a certain stage of development, once the vegetative organs are fully formed, the plant begins to initiate reproductive organs (sporangia, gametangia) where processes associated with plant reproduction take place.

Plant reproduction—that is, The ability to leave behind offspring—is one of the fundamental properties inherent in All living organisms. It sustains the continuity and succession of life on Earth. In plants, the reproduction process is highly complex and involves the alternation of sporophytic (asexual) and gametophytic (sexual) generations in The life cycle, meaning that developmental phases alternate through the strict succession of two reproductive processes: spore formation and the sexual process. The development of angiosperms is particularly intricate; they were termed 'winners in the Struggle for Existence' by the renowned Russian botanist and Moscow University professor M. I. Golenkin (1864–1941). A unique feature of angiosperm plants is The Emergence of a specialized reproductive organ—the flower—within which all reproduction-related processes occur. The flower also hosts the formation of seeds, through which flowering plants propagate.

Morphology can be understood in both a broad and a narrow sense. In the strict sense, morphology deals exclusively with the External structure of plants, whereas in the broad sense, this science also investigates internal structure, as well as the features of individual and historical (evolutionary) development. Thus, it encompasses The Study of both MACROSCOPIC AND MICROSCOPIC plant structures, alongside their developmental patterns. It is precisely in this comprehensive framework that the academic course 'Plant Morphology' is taught in biological higher education institutions, and it is from this perspective that this textbook has been written.

Morphology emerged as a scientific discipline from botany. Botany (from the Greek *botánicos*—relating to plants, *botane*—grass, herbs)—the science of plants—originated at the dawn of human history. Harsh living conditions, and above all the constant threat of starvation, compelled our distant ancestors to gather fruits, seeds, rhizomes, and other edible parts of wild plants. Thus, nomadic humans roaming vast open landscapes became acquainted with the beneficial and harmful properties of plants and learned to harness them for their own needs.

With the transition from a nomadic lifestyle to a settled one, agriculture began to take shape. In Ancient Egypt, several species of wheat, barley, and flax were cultivated as early as 3,000–4,000 BCE. Lentils, peas, broad beans, and many other valuable plants were brought into cultivation. Among fruit trees, grapes, date palms, figs (sycamores), and olives were widely grown. Agriculture was also highly developed in Ancient China. The famous Chinese scientist Shennong, in the manuscripts of *Shujing* (around 2,200 BCE), provided data on the properties and cultivation techniques of over 100 different plant species, including rice, sorghum, peas, beans, and cotton. Ancient China practiced a ritual sowing ceremony of cereal crops performed by the emperor himself. The Emperor of China, accompanied by a procession of lavishly dressed dignitaries (mandarins), walked behind a plow pulled by oxen, casting grains of wheat, barley,

rice, and soybean seeds into the plowed soil. This ritual was established by Emperor Shennong in 3,000 BCE.

In Ancient Rome, the cultivation of plants was considered not only vital but also a highly prestigious occupation. It was here, in Ancient Rome, that agricultural culture truly flourished. The Romans developed various soil fertilization Methods that remain in use today (such as applying wood ash and lime, and utilizing legumes as green manure).

In Ancient India, the study of medicinal plants became widespread. The oldest literary monument of Indo-European culture, the Yajur Veda (dating back over 2,000 years BCE), lists approximately 760 remedies, the majority of which are of plant origin. In Ancient Assyria and Babylon, as evidenced by cuneiform tablets, medicinal plants were extensively used to treat A wide variety of ailments.

A vast array of medicinal plants was employed in Ancient Greece. The ancient Greek physician Hippocrates recorded information in his writings on the healing properties of more than 200 species of local and foreign plants utilized in contemporary medicine.

Consequently, botany developed for a long time as an applied science pursued for purely utilitarian goals tied to agriculture and medicine.

The first attempt to synthesize numerous botanical facts was made by the great ancient Greek thinker and student of Aristotle (384–322 BCE), Theophrastus (372–287 BCE). Theophrastus was not merely a botanist; he was a philosopher, natural scientist, and polymath well-versed in many fields. With remarkable precision for his era, he formulated the objectives of botany as a science. In his landmark treatise *Enquiry into Plants*, he wrote: 'The differences between PLANTS AND THEIR general nature must be examined by investigating their parts, properties, distribution, and manner of life.' The works of Theophrastus laid the foundation for scientific botany, earning him the title of the 'father of botany' as aptly expressed by the outstanding Swedish botanist Carl Linnaeus (1707–1778).

The development of any science, including botany, cannot be viewed in isolation from the socio-economic and cultural-historical conditions of society. The early strides made by plant science during Antiquity stalled at the dawn of the Middle Ages, followed by a period of complete stagnation in botany linked to the economic and political decline of the Ancient World.

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The feudal system that succeeded the slave-owning society during the Early Middle Ages did little to foster scientific progress. It was not until the late 12th and early 13th centuries that noticeable shifts in the development of natural sciences began to emerge. This period saw the appearance of the scholarly works of Albertus Magnus, Bishop of Ratisbon (1193–1280), who authored seven books on plants. In them, from the perspective of his worldview, he attempted to explain plant life, the causes of winter dormancy, and plant nutrition.

In practice, however, for eighteen centuries (from the 3rd century BCE to the 16th century), the works of Theophrastus exerted an overwhelming influence on the development of botany. Throughout Antiquity and the Middle Ages, scientists advanced no further than him either in their understanding of plant evolutionary history or in the description of plant forms.

The revival of botany as a science began in the late 15th and early 16th centuries. This period was marked by the Age of Discovery. Driven by advancements in navigation, the Americas, Africa, and India were discovered, and Europeans became acquainted with many previously unseen plants and the spices derived from them, such as cinnamon, ginger, cloves, and black pepper.

The arrival of exotic plants in Europe compelled Europeans to seek methods for their preservation and study. The desire to preserve these specimens led to the invention of the herbarium method, proposed by the Italian botanist Luca Ghini (1490–1556). From 1528 to 1543, Luca Ghini served as a professor in Bologna and later in Pisa. Over time, Ghini's original herbarium collections were lost, but the herbaria of several of his students survived. From the 1630s and 1640s, driven by a surging interest in plant studies worldwide, herbarium collections began to be assembled by both professional botanists and nature enthusiasts. These herbaria varied greatly in scale. For instance, the Italian naturalist and multi-volume author on natural history Ulisse Aldrovandi (1522–1605) possessed 5,000 specimens in his herbarium; the French botanist and traveler Joseph Pitton de Tournefort (1656–1708) had 10,000; and the English naturalist John Ray (1627–1705) held 16,000. Carl Linnaeus considered his herbarium to be the largest, though records indicate it contained between 16,000 and 23,000 herbarium sheets, whereas the collection amassed largely through purchases from various travelers and scientists by Sir Hans Sloane (1660–1753), an English physician, collector, and president of the Royal Society, numbered 120,000 specimens.

In Russia, the earliest reliably documented plant collections that have survived to the present day (consisting of 100 herbarium specimens, housed in Saint Petersburg) were gathered by P. Areskin, personal physician to Peter the Great.

The oldest herbarium collections from Belarus, discovered by researchers from the Institute of Experimental Botany of the National Academy of Sciences of Belarus within the Vilnius University herbarium, date primarily to 1823–1826. One of the most extensive collections in this herbarium belonged to Stanisław Bonifacy Gorski, a florist-systematist, entomologist, and physician who served from 1824 to 1932 as director of the botanical garden at Wilno (now Vilnius) University, and subsequently as an adjunct professor of botany, pharmacy, and pharmacology at the Wilno Medico-Surgical Academy. The Herbarium of the Institute of Experimental Botany also preserves exsiccata sets published by R. H. Pabo and K. A. Czołowski (1854–1855), as well as N. V. Downar (1862).

The development of state herbaria engaged in research did not begin until the 19th century. According to the Index Herbariorum (1964, 1974), there are currently 23 major herbaria worldwide, each housing 2 million or more accessioned specimens. Medium-sized herbaria, holding at least 200,000 specimens, number around 170. Regional herbaria vary considerably in the size of their holdings.

The largest herbarium in Belarus is the Herbarium of the V. F. Kuprevich Institute of Experimental Botany at the National Academy of Sciences (MSK), which holds the official status of a National Treasure of the Republic of Belarus. It houses over 250,000 herbarium specimens, including 160,000 vascular plants, 37,000 bryophytes, 40,000 Lichens, 15,000 Fungi, and 125 species of algae. In addition, the Herbarium maintains modest paleobotanical and carpological collections.

Extensive long-term floristic studies conducted across the country under the guidance of Academician V. I. Parfenov, documented through substantial herbarium Materials, serve as a rich source of diverse data on plants, including morphological characteristics.

The second-largest national herbarium is that of the Belarusian State University (MSKU). It comprises approximately 75,000 specimens, featuring vascular plants of the Belarusian flora (around 50,000), a geographical herbarium (about 10,000), a mycological herbarium (primarily consisting of phytopathogenic micromycetes), and collections of lichens (4,000) and fungi (4,500). The herbarium also preserves a collection of plant seeds cultivated at the BSU Botanical Garden at various times.

While herbarium materials hold immense scientific value, the method of herbrizing has both advantages and limitations.

Because herbaria could not fully capture all the Structural Features of plants—particularly woody ones—botanical gardens began to be established in Europe to study living plants (for example, in Pisa, Italy, in 1543, and in Padua in 1545).

In Russia, the first botanical garden—an "apothecary garden"—was established in Moscow by decree of Peter the Great. A fragment of this historic garden has survived to the present day and is now part of the Moscow State University Botanical Garden.

The founder of the first botanical garden in Belarus was the French physician and botanist J. É. Gilibert (1741–1814), who arrived in Grodno in 1775 at the invitation of the St. Petersburg Academy of Sciences. Here he established a botanical garden (the exact date remains unknown) and founded a medical school where he taught until 1781. In 1782, Gilibert relocated to Vilna, taking the core collection of the botanical garden with him.

The Central Botanical Garden of the National Academy of Sciences in Minsk was founded in 1932 under the direction of Professor S. P. Melnik. Today, it ranks among the world's major botanical gardens, maintaining collections of approximately 10,000 plant species and cultivars grown in open-air plots and greenhouses. Since their inception, botanical gardens have played a vital role in plant Introduction research, maintained living plant collections, and served as key centers for science, education, and public outreach.

In 15th-century Europe, the demographic of those engaged in botany began to shift. While the science had previously been pursued primarily by monks—then the most educated members of society—the Renaissance saw a surge of interest among physicians and apothecaries. This period marked the emergence of specialized scholarly works known as herbals, which described Medicinal Plants and their Applications. The first herbal appeared in Europe between 1530 and 1536. Compiled by the German physician O. Brunfels (1470–1534) and titled Living Pictures of Plants, it effectively served as the first botanical atlas. In 1539, the German botanist H. Bock (1498–1544), known in the scientific community as Tragus, published the New Herbal. It featured descriptions and illustrations of 165 plant species, provided their local, Latin, and Greek names, and noted their flowering times, geographical distribution, and habitats. The New Herbal proved so popular that it went through 10 editions over the course of the 16th century.

As interest in botany flourished, an immense body of empirical data accumulated, becoming increasingly difficult to manage. In fact, until the mid-18th century, botany remained predominantly a descriptive and observational science. However, burdened by this vast accumulation of facts and scientific records yet continually enriched by new Research Methods and discoveries, the field gradually gave rise to a series of more specialized, focused, and concrete disciplines. Morphology is one such discipline.